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This  book  is  due  on  the  date  indicated 
below  and  is  subject  to  an  overdue  fine 
as  posted  at  the  Circulation  Desk. 


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DISEASE    IN    CAPTIVE   WILD 
MAMMALS    AND    BIRDS 

INCIDENCE,  DESCRIPTION,  COMPARISON 


BY 

HERBERT  FOX,  M.D. 

PATHOLOGIST  TO  THE  ZOOLOGICAL  SOCIETY  OF  PHILADELPHIA,  DIRECTOR  OF  THE  WILLIAM 
PEPPER  CLINICAL   LABORATORY,   UNIVERSITY  OF   PENNSYLVANIA 


WITH  A  FOREWORD  BY 
CHARLES  B.  PENROSE,  M.D. 

PRESIDENT  OF  THE  ZOOLOGICAL  SOCIETY  OF  PHILADELPHIA 


PHILADELPHIA,  LONDON  ^  CHICAGO 
J.  B.  LIPPINCOTT   COMPANY 


S" 


COPYRIGHT,    1923,    BY    HERBERT   FOX 


THIS  BOOK 
IS  DEDICATED  TO 

CHARLES  BINGHAM  PENROSE,  M.D.,  Ph.D.,  LL.D. 

IN   GRATEFUL  APPRECIATION  OF   HIS  FRIENDSHIP 
AND   OF   HIS   GUIDANCE  AND   ENCOURAGE- 
MENT IN  ALL  MATTERS  PERTAINING 
TO  THIS  LABORATORY. 


CONTENTS 


PAGE 

Foreword — Dr.  Charles  B.  Penrose 1 

SECTION 

I.    Introduction 17 

Zoological  Classification 

Numerical  List  of  Autopsies  Upon  Which   the  Work  is 
Based. 
II.    Diseases  of  the  Heart 48 

III.  Diseases  of  the  Blood  Vessels 66 

IV.  Diseases  of  the  Blood  and  Bone  Marrow 83 

V.    Diseases  of  the  Lymphatic  Tissues  Including  Spleen...  114 

VI.    Diseases  of  the  Respiratory  Tract 134 

VII.    Diseases  of  the  Alimentary  Tract. 

The  Tube  Proper 166 

The  Liver 222 

The  Pancreas 244 

The  Peritoneum 260 

VIII.    Diseases  of  the  Urinary  Tract 263 

The  Kidney 

IX.    Diseases  OF  the  Female  Reproductive  Organs 287 

X.    Diseases  of  the  Male  Reproductive  Organs 313 

XI,    Diseases  of  the  Ductless  Glands. 

The  Thyroid 316 

The  Thymus 336 

The  Suprarenal 336 

XII.    Diseases  of  the  Skeleton  and  its  Joints 343 

XIII.  Diseases  of  the  Central  Nervous  System  and  Special 

Senses , 372 

The  Eye 402 

XIV.  Constitutional  Diseases 410 

XV.    The  Relation  of   Diet  to  Disease   by  E.  P.   Corson- 
White,  M.D 415 

XVI.    Neoplasms 462 

XVII.    The  Infectious  Diseases. 

Tuberculosis 483 

Mycosis 558 

Streptothricosis 567 

Miscellaneous  Infections 596 

XVIII.    Animal  Parasites,  their  incidence  and  significance,  by 

F.  D.  Weidman,  M.D 614 


LIST  OF  TABLES 


PAGB 


1 .  The  incidence  of  degenerative  and  inflammatory  changes  in' the  heart.     5 1 

2.  The  incidence  of  hypertrophy  and  dilatation  of  the  heart  with  princi- 

pal associated  lesions 57 

3.  A  condensation  of  the  important  features  of  Tables  1  and  2 63 

4.  The  heart-to-body-weight  ratios 63 

5.  The  incidence  of  arteriosclerosis 71 

6.  The  differential  percentage  of  the  circulating  leucocytes  in  a  number 

of  different  animals 84 

7.  The  various  types  of  splenitis 126 

8.  The  incidence  of  the  various  types  of  bronchitis  and  of  pulmonary 

parasites 142 

9.  The  incidence  of  the  various  forms  of  pneumonia 150 

10.  An  analysis  of  cases  of  abscess  and  gangrene  of  the  lung 156 

1 1.  The  incidence  of  gastro-entero-colitis,  its  distribution  and  an  analysis 

of  the  causes 182 

12.  An  analysis  of  the  inflammations  of  the  alimentary  tract  in  Ungulata .   196 

13.  An   analysis   of   the    inflammations   of   the    alimentary,  tract    in 

Marsupialia 200 

14.  An  analysis  of  the  cases  of  pancreatitis,  the  associated  pathology 

and  the  details  of  the  anatomy  of  various  orders 252 

15.  The  weight  of  the  thyroid  body  in  relation  to  body  weight 318 

16.  The  incidence  of  the  various  enlargements  of  the  thyroid  body 326 

17.  The  incidence  of  cases  of  degenerative  bone  disease  in  the  various 

orders 357 

18.  The  weight  of  the  brain  and  the  relation  of  this  to  the  body  weights 

in  196  animals 388 

19.  An  analysis  of  pathological  conditions  in  terms  of  diet 423 

20.  An  analysis  of  the  diets  used  in  the  Philadelphia  Garden 426 

21.  The  incidence  of  tumors 464 

22.  An  analysis  of  the  breeding,  captivity  and  visceral  distribution  of 

neoplasms,  of  tumor-bearing  animals 466 

23.  An    analysis    of    the    incidence,    character    and     distribution    of 

tuberculosis 486 

24.  The  incidence  of  parasites  in  the  zoological  orders 630 

25.  The  incidence  of  heterakis  in  pheasants 635 

26.  The  incidence  of  various  parasitic  orders  and  families 636 

27.  The  visceral  distribution  of  parasites 639 

28.  The  occurrence  of  blood  parasites  in  zoological  classes 651 

vii 


FOREWORD 

BY 

CHARLES  B.  PENROSE 

The  work  on  which  this  book  is  based  was  begun  in 
November,  1901.  From  that  date,  systematic  autopsies 
were  made  on  animals  dying  in  the  Philadelphia  Zoo- 
logical Garden.  Previously  autopsies  had  been  made 
very  rarely  and  only  on  animals  of  especial  importance 
and  interest.  Pathological  conditions  were  occasionally 
noted  in  animals  subjected  to  anatomical  study.  The 
work  was  strictly  volunteer,  for  there  was  no  one  on  whom 
the  Society  had  the  right  to  call.  Dr.  Henrj'^  C.  Chapman, 
a  former  Director,  whose  interest  was  in  physiology  and 
comparative  anatomy,  made  nearly  all  the  autopsies  that 
were  made  before  the  beginning  of  the  present  work.  In 
the  annual  reports  from  the  foundation  of  the  Garden  in 
1872,  very  few  such  examinations  are  recorded.  In  some 
reports  there  are  lists  of  important  deaths,  but  no  record 
of  the  cause  of  death. 

This  state  of  things  was  not  peculiar  to  the  Philadel- 
phia Garden.  It  existed  in  every  zoological  garden  in  the 
world.  It  exists  in  most  of  them  today.  When  an  animal 
died  it  had  no  interest  or  value  except  for  its  hide  and 
bones.  Rare  specimens  were  sent  to  the  Academy  of 
Natural  Sciences  from  the  Philadelphia  Garden.  The 
great  majority,  however,  were  immediately  destroyed, 
and  thus  during  the  twenty-nine  years  from  the  founda- 
tion of  the  Garden,  preceding  this  work,  there  have  been 
lost  many  opportunities — some  of  which  will  never  recur 
— of  increasing  our  knowledge  of  pathology. 

This  book  gives  results  of  the  study  of  animals  that 
have  not  been  subjected  to  experimental  procedures  and 
conditions,  and  though  their  mode  of  life  has  not  been 

1 


2  FOREWORD 

that  strictly  natural  to  them,  yet  they  have  not  been  influ- 
enced by  any  of  the  artificial  procedures  of  the  laboratory 
which  are  usually  followed  in  investigations  on  animals. 
Though  the  object  of  the  work  was  the  pursuit  of  knowl- 
edge for  its  own  sake,  yet  results  of  practical  value  have 
followed:  hygiene  has  been  improved;  disinfection  has 
been  made  scientific ;  epidemics  have  been  arrested ;  some 
diseases,  notably  tuberculosis  in  monkeys,  and  spiro- 
pteriasis  in  parrots,  have  been  practically  eliminated. 

The  value  of  the  work  is  recognized  by  the  keepers. 
Their  interest  has  increased  and  their  morale  has 
improved.  The  frequent  deaths  in  a  large  collection  of 
animals  are  discouraging,  and  a  good  animal  man  must 
have  courage  and  optimism — traits  that  are  stimu- 
lated by  the  knowledge  that  every  animal  that  dies  is 
studied  to  determine  the  cause  of  death,  and  to  prevent 
its  recurrence. 

Some  results  of  this  study  are  to  be  considered  as 
*  *  raw  material, ' '  while  others  permit  cautious  or  tentative 
conclusions.  The  first  include — visceral  weights,  inci- 
dence of  certain  lesions  like  anemia  and  hepatic  cirrhoses, 
occurrence  of  calculi  and  observations  on  weak  hind 
quarters,  convulsions  and  constitutional  diseases.  The 
second  include — eradication  of  tuberculosis  and  spiro- 
pteriasis,  the  response  of  the  mammalian  and  avian  heart 
to  strain  and  disease  viruses,  the  origin  of  pneumonia  in 
birds,  the  genesis  of  bronchiectasis,  the  nature  of  osseous 
degeneration  with  relation  to  diet  and  alimentary  tract, 
the  comparative  anatomy,  physiology  and  pathology  of 
the  female  genitalia,  the  occurrence  of  lesions  in  the 
thyroid  comparable  to  those  in  man,  the  character  of 
kangaroo  disease. 

Great  praise  is  due  Dr.  Herbert  Fox  and  his  assist- 
ants Drs.  F.  D.  Weidman  and  E.  P.  Corson- White,  for 
the  splendid  scientific  spirit  and  thoroughness  mth  which 
they  have  done  this  work,  with  no  adequate  remuneration, 


FOREWORD  3 

except  the  intangible  reward  appreciated  only  by  the  real 
research  worker. 

It  is  a  mistake  to  think  that  all  animals  in  the  -wild 
state  are  healthy.  The  healthy  wild  animals  that  we  see 
are  the  survivors,  the  sick  and  the  weaklings  having  died. 
Undoubtedly  diseases  are  fewer  than  in  captive  animals ; 
but  on  the  other  hand  wild  animals  are  exposed  to  dangers 
to  which  captive  animals  are  not  exposed — such  as  lack  of 
care  in  sickness  and  old  age,  starvation,  and  attacks  of 
enemies  that  feed  on  them.  The  maximum  longevities  in 
some  species  are  probably  in  captive  animals.  It  is  doubt- 
ful if  a  wild  animal  ever  reaches  the  greatest  age 
possible  to  it. 

Many  kinds  of  parasites  have  been  found  in  wild 
animals  captured  only  a  short  time  before  arriving  at  the 
Garden,  so  that  they  must  have  existed  in  the  wild.  Thir- 
teen wild  cats  received  from  South  Carolina  within  a  few 
days  after  capture  were  infested  with  intestinal,  muscu- 
lar, and  pulmonary  parasites.  Several  years  ago  there 
were  received  from  the  Island  of  Chincoteague,  off  the 
coast  of  Virginia,  a  number  of  native  birds  recently  cap- 
tured, all  of  which  had  mould  disease  of  the  air  sacs. 
This  happened  on  several  occasions,  so  that  it  became 
necessary  to  reject  all  birds  from  this  locality.  Among 
the  birds  were  meadow  larks  and  cardinals. 

Hunters  not  infrequently  kill  animals  with  disease, 
and  if  more  hunters  knew  enough  to  recognize  the  disease 
there  would  be  more  recorded  cases.  Sick  animals  are  not 
as  Ukely  to  be  killed  by  the  hunter  as  healthy  ones,  as  sick 
animals  seek  seclusion  and  do  not  move  about.  In  many 
localities  of  the  United  States  white-tailed  deer  are 
infested  with  the  liver-fluke.  Trout  and  other  fish  in 
remote  mountain  streams  are  found  with  tapeworms. 
Round  worms  and  other  parasites  infest  the  white 
rhinoceros  in  the  African  forest. 


4  FOREWORD 

In  many  instances  the  parasites  and  the  host,  when  in 
health,  get  on  very  .well  together ;  but  when  the  host  weak- 
ens the  parasites  may  give  him  the  finishing  stroke. 

Though  some  morbid  conditions  described  in  this  book 
are  peculiar  to  capti^dty,  yet  it  seems  reasonable  to 
assume  that  many  of  the  diseases  found  in  captive  ani- 
mals occur  also  in  the  wild.  Some  of  the  extinct  races  of 
animals  may  have  been  wiped  out  by  disease. 

Captivity  causes  numerous  physical  and  mental  dis- 
arrangements. Unaccustomed,  unnatural  and  unvaried 
food,  change  of  climate  and  environment,  physical  and 
mental  degeneration  from  disuse  of  muscle  and  brain, 
fear,  ennui,  nostalgia,  lack  of  the  exhilaration  of  chasing 
and  being  chased,  unsatisfied  sexual  feeling — all  react 
harmfully  on  the  captive. 

No  captive  animals  get  their  natural  food ;  and  though 
some,  like  the  carnivora,  may  get  approximately  their 
natural  food,  yet  they  do  not  get  it  in  the  natural  way. 
They  have  but  little  variety  and  may  miss  elements  impor- 
tant to  their  well-being.  They  get  only  certain  cuts — 
muscle  and  bone ;  they  do  not  get  blood,  guts  and  glands. 
The  lion 's  meat  is  handed  to  him.  He  does  not  tear  down 
his  prey ;  and  one  result  is  that  the  jaws  of  the  captive- 
bred  lion  differ  decidedly  in  shape  from  the  jaws  of  the 
T\^ld-bred  lion.  He  gets  his  food  regularly,  with  awful 
monotony — twelve  pounds  of  meat  at  3.30  p.m.  da}'^  after 
day;  there  is  no  alternation  of  feasts  and  fasts,  with 
consequent  change  in  the  balance  of  the  body  reserves. 

Some  animals,  such  as  caribou,  the  moose,  the  black 
cock,  the  hoatzin,  the  koala,  do  not  long  survive  captivity, 
because  it  is  impracticable  to  obtain  their  natural  food. 
Gastrointestinal  disease  is  the  commonest  disease  of 
wild  animals.  Twenty-five  per  cent,  of  sick  humans  in 
civilized  communities  also  suffer  from  it.  On  the  other 
hand,  many  animals  become  accustomed  to  the  new  diet 
and  thrive  on  it. 


FOREWORD  5 

Animals  also  often  thrive  in  a  climate  very  different 
from  that  of  their  natural  habitat.  Some  animals  from 
warai  comitries,  though  kept  outside  without  any  artificial 
heat,  get  on  very  well  during  the  severe  weather  of  Phila- 
delphia winters,  when  the  temperature  often  drops  to 
zero  Fahrenheit.  For  the  past  two  years  it  has  been  the 
custom  to  put  outdoors  all  monkeys — of  value — that  are 
suspected  of  having  tuberculosis.  The  cheap  monkeys  are 
killed.  The  outdoor  monkeys  are  kept  in  fair  sized  cages — 
five  feet  square  and  six  feet  high — usually  singly,  though 
sometimes  two  are  together.  The  vervet,  the  grivet,  the 
leonine  macaque,  the  yellow  baboon,  the  Hainan  gibbon 
and  many  others,  have  not  only  survived  the  climatic  con- 
ditions but  have  not  succumbed  to  tuberculosis.  South 
American  monkeys  do  not  stand  cold  as  well  as  Old  World 
monkeys,  and  cannot  be  kept  out  in  severe  winter  weather. 

The  monkeys  that  are  kept  indoors  during  winter  have 
free  access  to  the  outside  through  swinging  doors  and 
they  often  go  out  voluntarily  in  the  coldest  weather.  The 
same  is  true  of  other  animals.  A  note  made  several  years 
ago  (February  12, 1914)  states :  ''Temperature  last  night 
was  1°  F.  At  2  P.M.  outside  temperature  13°  F.,  in  car- 
nivora  house  25°  F.,  elephant  house  38°  F.,  giraffe  house 
39°  F.,  monkey  house  42°  F.  Two  Bengal  tigers  were 
voluntarily  outside.  Monkeys  that  have  been  out  all 
winter  are :  two  Barbary  apes,  Hainan  gibbon,  lion-tailed 
macaque,  yellow  baboon,  grivet  monkey,  pig-tailed 
macaque,  and  eleven  rhesus  macaques.  All  the  animals 
appeared  comfortable." 

The  physical  condition  of  the  animal  and  the  kind  of 
cold — damp  or  dry — have  much  to  do  with  its  ability  to 
stand  low  temperatures.  Monkeys  have  passed  through 
very  severe  winters  without  damage,  and  subsequently  in 
a  less  severe  winter  have  had  frozen  fingers,  toes  or  tail. 

The  size  of  the  cage  or  pen  has  not  as  much  effect  upon 
the  well-being  of  the  animal  as  might  be  expected.  Rep- 
tiles, birds  and  mammals  do  as  well  in  cages  and  pens  of 


6  FOREWORD 

medium  size  as  in  very  large  ones.  A  deer  or  antelope  in 
a  large  enclosure  does  not  use  all  its  domain.  It  usually 
has  a  favorite  corner  near  the  food,  water  and  shelter, 
and  stays  there.  Nor  has  a  large  enclosure  been  found 
perceptibly  to  diminish  mortality  from  cage-mates.  The 
stronger  will  follow  the  weaker  until  he  gets  him,  no 
matter  what  the  enclosure.  Even  in  the  large  flying  cage 
for  birds  it  is  necessary  to  keep  out  those  of  a  scrappy 
disposition.  From  the  point  of  view  of  the  public  small 
enclosures  are  more  satisfactory. 

Mental  disease  in  captive  animals  offers  a  fasci- 
nating field  for  study — now  chiefly  speculative.  Many 
conditions  are  present  to  produce  it — all  the  con- 
ditions that  cause  prison  psychoses  in  man.  And 
many  if  not  all  the  insane  and  perverted  acts 
of  animals  have  their  counterpart  in  the  human. 
Masturbation  is  very  common  in  mammals.  Eating  of 
their  own  feces,  coprophagy,  is  not  infrequent,  and  is 
especially  notable  in  one  of  the  highest  types — the  Chim- 
panzee. Perhaps  occasionally  coprophagy  may  be  due  to 
some  defect  in  diet.  There  is  no  instinctive  disgust  at 
excrement  in  the  lower  animals  any  more  than  there  is  in 
the  uneducated  child.  Nevertheless,  eating  feces  cannot 
be  healthful,  and  probably  does  not  occur  in  nature ;  and 
occurs'  only  in  the  human  with  mental  disease. 

Some  of  the  insane  acts  of  animals  if  prevalent  in  the 
wild  would  probably  cause  the  extinction  of  the  race. 
Such  are  killing  of  the  young  by  the  father  and  by  the 
mother ;  killing  of  the  female  by  the  male,  usually  during 
rutting  time,  in  some  instances  reminding  one  of  Sadism 
in  man.  This  kind  of  sexual  killing  does  not  often  occur 
in  the  wild.  The  female  has  a  better  chance  to  escape, 
and  the  male  probably  does  not  feel  so  inclined  to  damage 
her  when  he  chooses  her  himself  as  when  she  is  chosen  for 
him  by  his  gaoler.  When  the  mother  devours  her  young 
it  is  usually  shortly  after  birth.  I  have,  however,  the 
record  of  a  Jungle  cat  {Felis  chaus)  who  ate  two  of  her 


FOREWORD  7 

kittens  when  they  were  seven  weeks  old.  Some  of  the 
domestic  animals  devour  their  young ;  the  sow  often  does 
it,  and  occasionally  the  bitch. 

Sucking,  gnawing  and  eating  parts  of  itself  or  of  a 
cage-mate  is  not  uncommon  in  a  variety  of  animals.  Bears 
lick  their  paws  until  they  are  sore ;  a  monkey  may  gradu- 
ally gnaw  away  its  tail  from  the  tip  to  the  body ;  an  ocelot 
(Felis  chihigonazon)  bit  open  his  scrotum  and  devoured 
his  testicle;  a  Tasmanian  devil  (Sarcophilus  ur sinus)  bit 
off  one  of  his  front  feet  at  the  wrist;  a  monkey  may 
gradually  gnaw  away  its  fingers;  and  numerous  other 
self-inflicted  mutilations  occur.  Often  there  is  a  local 
irritating  cause,  as  skin  disease,  lice,  or  freezing.  But 
in  many  cases  no  local  cause  can  be  found,  as  no  local 
cause  can  be  found  for  thumb-sucking  or  nail-biting  in 
the  human. 

The  surgery  done  by  monkeys  on  their  frozen  fingers 
and  toes  is  interesting.  After  the  flesh  has  sloughed  the 
monkey  bites  off  the  protruding  phalanges,  apparently 
without  pain,  so  that  satisfactory  well-covered  stumps 
are  made. 

Animals  often  mutilate  their  cage-mates  in  an 
amicable  way  as  distinguished  from  fighting.  A  bear  may 
lick  its  mate's  ears  until  the  hair  and  skin  are  gone.  A 
monkey  may  eat  its  mate's  tail  or  patches  of  its  skin,  the 
victim  lying  placidly  while  the  process  goes  on.  Many 
animals  are  addicted  to  perverted  acts  on  their  own  or 
their  mate's  sexual  organs. 

It  is  probable  that  the  phenomena  just  mentioned  are 
due  to  confinement,  idleness  and  ennui,  and  that  they  do 
not  often  happen  in  nature  where  an  animal  is  kept  busy 
seeking  his  food,  fighting  and  avoiding  his  enemies, 
attending  to  his  mate  or  mates,  and  meeting  the  various 
vicissitudes  of  his  environment. 

It  should  be  remembered,  however,  that  the  members 
of  a  wild  species  vary  in  intelligence  and  temperament,  as 
humans   do.     There   are  morons   and   perverts   among 


8  FOREWORD 

animals  in  the  \\'ild ;  but  not  being  coddled  by  the  normal 
members  of  the  species,  they  have  a  poorer  chance  than 
has  the  subnormal  human  of  surviving  and  of  transmit- 
ting their  peculiarities. 

Fear,  ennui,  loneliness  and  nostalgia,  by  affecting  the 
minds  of  captive  animals,  react  on  their  physical  condi- 
tion. Some  animals  have  the  fear  of  man  bred  in  them. 
The  young  often  show  it  from  time  of  birth.  This  is 
especially  common  in  animals  that  have  survived  for  gen- 
erations in  proximity  to  man.  It  is  one  reason  for  their 
survival.  The  mother  and  father  may  have  become  tame 
and  gentle  in  captivity  and  yet  the  young  one  may  be  a 
wild  thing  from  birth.  Such  fear  is  sometimes  uncontrol- 
lable, an  apparently  slight  cause  making  the  animal  dash 
itself  against  the  fence  of  the  enclosure.  It  is  not  mere 
speculation  to  discuss  the  physical  effects  of  the  emotions 
on  the  animal  body.  It  has  been  shown  that  fear,  anger, 
and  grief  bring  about  distinct  measurable  physical 
changes.  Dr.  Corson- White  has  found  that  the  red  cor- 
puscles are  increased  by  over  two  million  per  cubic 
centimetre  in  the  blood  of  a  cat  frightened  by  a  dog  bark- 
ing at  it.    The  amount  of  blood  sugar  is  also  increased. 

Such  observations  are  suggestive  in  a  consideration 
of  the  changes  that  may  occur  in  a  captive  animal  sub- 
jected to  acute  and  chronic  fear. 

The  monotony  of  a  captive  animal 's  life  is  broken  only 
by  feeding,  the  sight  and  sounds  of  others  in  the  same 
building  or  nearby,  and  by  visitors.  Many  animals  show 
their  appreciation  and  pleasure  when  visitors  approach, 
and  some  of  the  more  intelligent  animals,  bears  and 
monkeys  and  some  birds, ''  show  off  "  apparently  to  keep 
the  visitors  there.  When  there  is  nothing  doing,  some 
stand  swaying  their  heads,  like  a  weaving  horse,  or  pacing 
the  cage,  inanely  tagging  at  each  turn  the  side  of  the  cage 
with  the  head  or  other  part  of  tlie  body — often  so  persist- 
ently that  a  sore  is  produced.  Nearly  all  animals  are 
social  and  suffer  from  loneliness  when  kept  by  themselves. 


FOREWORD 

This  is  true  even  of  the  lowly  forms.  The  keeper  of  the 
reptile  house  reported  that  a  giant  tortoise  became 
despondent  and  refused  to  eat  when  his  companion,  a 
leopard  tortoise,  was  taken  from  him,  and  that  he  braced 
up  as  soon  as  the  leopard  tortoise  was  returned.  It  is  not 
necessary  that  the  companion  be  of  the  same  species,  or 
even  of  the  same  family.  A  lion  or  a  tiger  may  be  satis- 
fied with  a  little  dog  for  a  companion,  and  there  was  an 
African  rhinoceros  at  the  Philadelphia  harden  that  was 
very  discontented  and  unhappy  when  alone  and  became 
perfectly  satisfied  when  she  was  given  a  domestic  goat  as 
a  cage-mate;  and  the  huge  rhinoceros  stood  for  a  good 
deal  of  butting  and  bullying  from  the  goat  without 
retaliating.  A  sympathetic  keeper  may  do  much  to  relieve 
the  loneliness  of  the  animals  in  his  care. 

Nostalgia,  or  homesickness,  has  been  felt  by  all  men. 
Some  have  died  of  it.  The  tradition  among  writers  is 
that  it  affects  young  people  and  those  who  have  been 
living  nearest  to  a  state  of  nature.  In  this  country  the 
American  Indian  and  the  negro  are  affected  more  than 
the  whites.  Much  was  written  of  it  after  the  Franco- 
Prussian  War  and  the  American  Civil  War.  It  is  a  real 
condition,  capable  in  extreme  cases  of  causing  death  and 
of  so  weakening  the  sufferer  as  to  make  him  more  suscep- 
tible to  the  invasion  of  other  diseases.  At  the  present  day 
we  hear  less  of  it  among  civilized  people  than  formerly, 
perhaps  because  the  conception  of  home  has  been  broad- 
ened by  modem  methods  of  intercommunication.  The 
wild  animal's  conception  of  home  is  narrow;  he  comes 
directly  from  it  into  an  environment  where  he  may  see 
many  other  animals,  but  not  one  of  his  own  kind.  Pre- 
disposing causes  of  nostalgia  are  stronger  with  him  than 
with  the  human.  That  home  means  a  great  deal  to 
animals  is  shown  by  the  migration  of  birds — the  return  of 
the  carrier  pigeon,  and  of  the  lost  dog,  and  of  the  swallow, 
w^hich  returns  every  year  to  the  same  nesting  place. 

2 


10  FOREWORD 

All  animals  long  for  the  things  of  nature — open  air, 
earth,  grass  and  water.  They  are  thrilled  when  their  feet 
touch  the  sod.  Even  the  hippopotamus  gambols  when  he 
leaves  his  concrete  house  and  his  feet  touch  earth 
and  grass. 

The  face  and  carriage  of  many  animals  cannot  express 
feelings  as  in  the  human,  though  it  is  not  unreasonable  to 
assume  that  animals  may  indicate  feelings  by  expression 
understood  by  their  mates,  though  not  understood  or  even 
noticed  by  man.  When  they  can  express  it  in  a  human 
way  their  feelings  may  be  read.  The  dejection  of  nos- 
talgia is  especially  shown  by  anthropoid  apes.  Gorillas 
have  been  noted  for  it  from  the  earliest  writers.  The 
orang  is  prone  to  it,  shows  it  by  his  attitude  and  expres- 
sion, and  sometimes  dies  of  it. 

It  is  stated  in  A  Handbook  of  the  Management  of 
Animals  in  Captivity  in  Lower  Bengal,  p.  130,  that 
elephants  have  been  observed  to  shed  tears  abundantly  if 
forced  to  leave  their  old  home  and  surroundings.  How 
much  other  animals  who  cannot  express  homesickness 
may  feel  it,  and  how  often  it  is  a  cause  of  unhappiness, 
depression  and  predisposition  to  disease,  it  is  impossible 
to  say. 

Most  wild  animals  in  captivity  are  sterile.  The  reason 
is  not  known.  It  shows  the  profound  effect  of  captivity. 
It  would  be  diflScult  to  determine  whether  the  sterility 
of  a  mating  is  the  fault  of  the  male  or  the  female. 

There  is  no  apparent  rule  for  sterility.  Some  families 
are  always  sterile  in  captivity,  others  are  fertile,  even 
with  very  unfavorable  surroundings.  The  deer,  horses, 
hippopotamuses,  pigs,  goats,  sheep  and  oxen,  are  good 
breeders;  while  antelopes,  rhinoceroses,  giraffes,  ele- 
phants, are  poor  breeders.  Some  members  of  a  family 
may  be  good,  others  poor.  The  lion  and  puma  breed  fairly 
well ;  the  tiger,  leopard  and  jaguar,  very  poorly.  Bears 
breed  well,  but  the  mother  usually  destroys  her  young. 


FOREWORD  11 

We  cannot  mate  wild  animals  and  birds  simply  by 
putting  males  and  females  together  in  the  same  cage. 
Domestic  mammals  and  birds  usually  mate  under  such 
conditions,  but  wild  ones  often  refuse.  Many  mammals 
and  probably  all  birds  that  are  not  polyandrous  or 
polygamous  reserve  the  right  to  select  their  mates,  and  if 
the  sexes  are  put  together  by  man  they  may  view  each 
other  with  indifference  or  with  animosity.  There  are 
many  males  and  females  of  the  same  species  of  mammals 
at  the  Zoological  Garden  that  will  not  consent  to  live 
together.  A  male  monkey  in  a  cage  with  several  females 
will  very  often  select  one  female  for  his  mate  and  will 
have  nothing  to  do  with  the  others. 

Among  monkeys  fertility  varies  greatly.  It  is  not 
practical  to  determine  the  ratio  among  the  various  kinds, 
as  some  kinds  are  much  commoner  in  zoological  collec- 
tions than  others.  I  think  that  in  general  the  Old  World 
monkeys  (Cercopithecidce)  are  better  breeders  in  cap- 
tivity than  those  of  the  New  World  (Cebidce).  The 
anthropoid  apes  are  very  poor  breeders  indeed;  of  the 
many  gibbons,  orangs,  and  chimpanzees,  that  for  years 
have  been  captive  in  Europe  and  America,  it  is  probable 
that  only  the  chimpanzee  has  bred,  and  that  very  rarely. 

Refusal  to  mate,  sterility,  infanticide  by  father  and 
mother,  and  sexual  killing  keep  down  reproduction  in 
zoological  gardens;  and  the  number  of  young  ones  is  a 
good  indication  of  the  character  of  a  garden  and  of  the 
provisions  made  for  the  happiness,  comfort  and  health  of 
the  animals. 

With  the  birds  in  a  zoological  garden  the  conditions 
for  nesting  and  laying  are  not  good.  Caged  birds  have  no 
material  for  a  nest,  no  privacy  and  rarely  lay  an  egg. 
The  outdoor  water  birds  and  the  outdoor  upland  birds 
with  natural  surroundings,  with  secluded  retreats,  lay 
and  hatch  very  well.  Birds  like  mammals  apparently 
are  indifferent  to  publicity  when  copulating,  but  seek 


12  FOREWORD 

seclusion  for  laying  and  nesting — from  maternal  fear  for 
the  safety  of  the  young. 

The  sexual  instinct  in  indoor  caged  birds  in  a  zoologi- 
cal garden  is  dormant.  Very  few  copulate  and  very  few 
lay  eggs;  pigeons  and  Mexican  conures  {Conurus  liolo- 
chlorus)  are  exceptions  to  this  general  rule. 

Probably  because  the  sexual  instinct  is  dormant  the 
males  never  fight  over  the  females  among  perching  birds, 
and  veiy  rarely  kill  each  other.  In  some  species  of 
finches,  however,  as  the  chestnut-eared  {gn.  Amadina), 
the  females  fight  among  themselves  if  there  are  not 
enough  males. 

Ovoviviparous  reptiles  breed  more  often  in  captivity 
than  one  would  expect;  and  egg-laying  snakes  often  lay 
eggs,  which  of  course  are  only  hatched  artificially. 

Birds  suffer  less  than  mammals  from  the  psychologi- 
cal effect  of  captivity.  The  mental  development  of  a  bird 
is  much  lower.  With  few  exceptions,  like  the  ruffed 
grouse,  the  bird  accepts  captivity  easily  and  becomes 
tame,  though  he  will  not  stand  being  touched.  He  views 
his  keeper  and  visitors  with  indifference  or  friendliness. 
If  a  bird  house  is  bright,  cheerful  and  sunny,  all  the 
inmates  thrive  and  appear  to  be  happy.  These  conditions 
undoubtedly  affect  the  health  of  the  bird,  as  is  evidenced 
by  their  plumage;  bright  colors  that  are  lost  in  a  dark 
and  gloomy  house  are  retained  and  developed  when  the 
house  is  cheerful  and  sunny.  There  are  some  birds,  how- 
ever, that  never  retain  their  colors  in  the  captivity  of  a 
zoological  garden.  Among  them  are  the  scarlet  ibis,  the 
American  flamingo,  and  the  roseate  spoonbill.  It  has 
been  suggested  that  the  loss  of  color  is  due  to  the  lack  of 
something  in  the  diet,  mineral  or  organic,  that  the  bird 
gets  in  its  natural  habitat.  Tame  scarlet  ibises  living 
with  the  chickens  about  the  dwellings  of  natives  in 
Venezuela  retain  their  brilliant  color.  The  material  of 
the  beautiful  red  color  on  the  under  surface  of 
the  wings  of  the  touracou  contains  copper,  yet  these 


FOREWORD  13 

birds  retain  this  color  very  well  in  captivity,  even  after 
several  moultings. 

The  source  of  the  copper  has  long  been  uncertain.  In 
nature  the  birds  are  fruit-eaters,  and  their  diet  in  cap- 
tivity consists  of  bone  meal,  zweibach,  corn  meal,  wliite 
potatoes,  eggs  and  carrots — foods  that  are  usually 
assumed  to  contain  no  copper;  and  no  copper  utensils  are 
used  in  the  preparation  of  the  food.  Dr.  John  Marshall, 
however,  writes  me  that  all  the  common  cereals  contain 
minute  quantities  of  copper;  and  Dr.  Leon  A.  Ryan, 
University  of  Pennsylvania  Medical  Bulletin,  June,  1907, 
states  that  copper  may  be  found  in  animal  tissues.  Dr.  E. 
P.  Corson- White  has  found  traces  of  copper  in  the  bone 
meal  used  at  the  Philadelphia  Zoological  Garden.  The 
copper  in  the  red  color  of  the  touracou's  wing  therefore 
comes  from  the  food. 

It  is  probable  that  a  bright  and  cheerful  bird  house 
does  not  influence  the  color  of  birds  by  the  direct  action 
of  light  on  the  color  as  much  as  indirectly  by  improv- 
ing the  health  and  spirits  of  the  birds.  Coloration 
in  birds  is  a  very  complicated  proposition.  It  depends 
upon  age,  sex,  season,  health,  light,  heat,  moisture,  mode 
of  life,  and  food.  No  one  bird  house  can  combine  all  the 
conditions  necessary  for  the  retention  of  natural  colors 
in  every  species.  The  desert  species  from  a  habitat  of 
intense  light  and  dryness  require  for  their  color  a  dif- 
ferent environment  from  the  forest  species,  from  a 
habitat  of  shade  and  moisture.  The  suppression  of 
sexual  feeling  in  captive  birds  may  influence  color.  In 
nature  the  finest  colors  are  attained  by  mating  birds. 

It  may  be  said  that  all  animals — except  those  of  noc- 
turnal habit — have  a  feeling  of  joy  and  well-being  in  fine 
-weather  and  bright  surroundings  that  reacts  favorably  on 
the  general  health. 

The  variability  of  the  breeding  period  induced  by 
captivity  in  many  animals  may  be  mentioned  with 
sterility.    It  was  sho\\Ti  some  years  ago  in  the  Philadel- 


14  FOREWORD 

phia  Garden  by  the  European  brown  bear  which  in  one 
year  gave  birth  on  January  16th,  and  in  the  following 
year  on  July  25th.  It  is  another  evidence  of  the  profound 
effect  of  captivity  on  the  captive  animal.  I  know  of  no 
observations  of  the  effect  of  captivity  on  the  period 
of  gestation. 

There  is  considerable  mortality  among  captive 
animals  from  killing  of  cage-mates.  I  do  not  refer  to 
sexual  kilhng,  already  mentioned,  or  to  fights  over  a 
female.  Often  males,  with  no  females  near,  cannot  be 
kept  together ;  probably  sexual  jealousy  is  at  the  bottom 
of  it.  Antelope  and  deer  are  especially  inclined  to  scrap. 
Even  a  large  enclosure  will  not  save  the  weaker  male ;  the 
stronger  follows  him  with  horrible  persistency,  some- 
times for  days,  around  and  around  the  enclosure,  often 
at  a  walk,  but  always  on  the  offensive,  at  least  during  the 
day;  until,  careless  from  weariness,  the  weaker  is  caught 
unawares  and  finished  by  a  horn-thrust  in  the  side. 

Both  birds  and  mammals  often  kill  their  mates  when 
the  mate  is  sick,  or  ^'  down  "  from  injury  or  disease.  All 
animals  hate  sickness  and  death,  and  show  their  dislike 
by  attacking  or  shunning  it.  Birds  may  get  on  happily 
together  for  months  until  one  becomes  sick,  and  as  he 
crouches  in  a  corner  with  ruffled  feathers  the  others  pick 
on  him  and  finish  him.  The  same  is  true  of  mammals,  the 
sick  one  being  horned  or  tramped  to  death  by  the  mate 
with  whom  he  had  formerly  been  on  most  friendly  terms. 
The  keeper  often  reports  an  animal  "killed  by  its  mate," 
whereas  the  mate  has  only  given  the  coup  de  grace. 

This  brutality  is  not  universal.  Rarely  a  parrakeet 
will  stand  guard  over  his  sick  and  dying  mate;  and  we 
have  seen  a  ratel — of  a  ferocious  family — stand  guard 
over  and  resist  the  removal  of  his  sick  companion. 

The  diagnosis  of  disease  in  wild  animals  is  unsatis- 
factory ;  usually  impossible ;  clinical  study  as  we  know  it 
in  the  human  is  impossible.  "We  know  that  the  animal  is 
sick,  but  not  why.   A  certain  group  of  symptoms  accom- 


FOREWORD  15 

panies  all  diseases — dull,  rough  coat  or  feathers,  refusal 
to  eat,  weakness  in  the  hind  quarters,  and  finally  getting 
down.  They  rarely  show  symptoms  of  pain — or  at  least 
we  cannot  read  the  symptoms.  The  pain  of  acute  pan- 
creatitis in  man  is  violent,  yet  many  animals  die  with  it 
and  we  cannot  tell  that  they  suffer.  Animals  do  not  suffer 
as  much  as  the  human,  and  they  stand  the  ravages  of 
disease  better  than  the  human.  At  autopsy  we  often 
wonder  how  the  animal  could  have  lived  with  the  condi- 
tions that  are  found.  A  monkey  may  be  apparently  well 
until  a  few  days  before  his  death,  though  his  lungs  and 
abdominal  organs  may  be  a  mass  of  tubercle.  A  small 
red  howler  monkey  {Alonata  seniciilus)  was  in  apparent 
good  health,  playful  and  lively  until  twenty-four  hours 
before  his  death  from  acute  pancreatitis,  though  liis 
stomach  and  intestines  contained  fifty-one  nematode 
worms,  some  of  which  were  eight  inches  in  length. 

As  diagnosis  is  unsatisfactory,  so  is  treatment. 
Usually  all  we  can  do  is  to  treat  symptoms ;  and  by  the 
time  disease  has  advanced  to  the  point  of  becoming  exter- 
nally noticeable,  it  has  usually  gone  beyond  the  reach  of 
medical  treatment.  It  must  also  be  remembered  that 
drugs  vary  very  much  in  their  action  in  different  families 
of  animals.  Nux  vomica  will  not  kill  the  gallinaceous 
birds  of  North  America,  and  Tenant  says  that  in  Ceylon 
the  hornbill  feeds  on  the  fruit  of  strychnos  nux  vomica. 
The  pigeon  is  immune  to  opium.  The  Felidce  are  said  to 
be  unusually  susceptible  to  carbolic  acid ;  veratrum  viride 
is  harmless  to  sheep  and  elk,  but  poisonous  to  the  horse ; 
dogs  can  take  with  impunity  large  quantities  of  cyanide 
of  potassium.  These  statements  are  true  when  the  drugs 
are  administered  by  mouth— the  usual  way  of  giving  them 
to  wild  animals.  The  action  may  be  different  if  the 
drugs  are  administered  intravenously  or  subcutaneously. 
Variations  in  effect  when  they  are  administered  by  mouth 
are  often  due  to  chemical  variations  in  the  digestive  secre- 
tions.   It  is  probable  that  the  action  of  cyanide  of  potash 


16  FOREWORD 

on  dogs  depends  on  the  amount  of  hydrochloric  acid  in 
the  digestive  tract. 

When  worms  or  their  eggs  are  found  in  the  stools 
vermifuges  are  used,  and  with  some  animals  especially 
liable  to  infestation  by  intestinal  worms,  periodic  doses  of 
vermifuges  are  given  as  a  prophylactic.  Turpentine  is 
given  to  the  zebra  at  fixed  intervals  for  the  round  worm ; 
santonin,  male  fern  and  areca  nut  to  the  carnivora  for  the 
various  worms  that  are  so  common  in  the  intestinal  tracts 
of  these  animals. 

The  work  of  the  Laboratory  of  Pathology  is  throwing 
light  on  the  subject  of  diagnosis,  and  though  from  the 
character  of  the  clinical  material  diagnosis  can  never  be 
as  satisfactory  as  in  the  human,  yet  we  may  fairly  hope 
for  improvement.  Prophylaxis  is  our  chief  reliance,  and 
always  will  be. 


DISEASE   IN    CAPTIVE   WILD 
MAMMALS  AND  BIRDS 

SECTION  I 
INTRODUCTION 

"  We  have  also  parks  and  enclosures  of  all  sorts,  of 
beasts  and  birds;  which  we  use  not  only  for  view  and 
rareness,  but  likewise  for  dissections  and  trials,  that 
thereby  we  may  take  light  Avhat  may  be  wrought  upon  the 
body  of  man." 

The  purpose  of  a  menagerie  under  the  auspices  of  a 
zoological  society  can  scarcely  be  put  into  better  words 
than  those  found  in  this  quotation  from  Sir  Francis 
Bacon's  New  Atlantis.  Apt  as  this  description  of  the 
mythical  island's  collection  may  be,  it  is  but  a  reflection 
of  the  teachings  of  Plato 's  original  legend  of  a  perfected 
community,  and  the  practical  applications  of  these  teach- 
ings by  Aristotle  in  his  Anatomy  and  Physiology  of 
Animals.  The  history  of  human  study  shows  a  constant 
investigation  of  lower  forms  of  life,  ever  broadening  in 
its  scope,  ever  more  satisfying  in  its  explanation  of 
biologic  problems  and  ever  increasing  in  value  from  an 
economic  standpoint. 

If,  however,  all  animals  are  to  be  subjected  to  ''dis- 
sections and  trials"  there  inevitably  will  come  under 
observation  many  specimens  presenting  variations  from 
the  accepted  mean  or  standard  or  even  from  an  average 
for  their  kind  and  therefore  approaching  what  may  be 
called  pathological. 

The  desire  to  explain  the  abnormal  has  had  the  effect, 
during  the  half  century  since  Virchow  defined  cellular 
pathology  and  Darwin  systematized  the  world's  knowl- 
edge of  comparative  biology,  of  directing  attention  to 
comparative  pathology  and  of  stimulating  the  study  of 

17 


18     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

veterinary  medicine.  Moreover  in  the  past  twenty-five 
years  much  work  has  been  done  and  many  isolated  pub- 
lications have  appeared  upon  the  diseases  of  wild  animals, 
notably  Bland-Sutton's  work,  Evolution  and  Disease 
(1895),  a  thoroughly  scientific  and  most  charmingly  writ- 
ten book,  but  rather  elementary  in  its  approach  of  the 
subject  of  pathology,  and  Wood  Hutcliinson's  Diseases  of 
A7iimals,  a  more  or  less  popularly  presented  treatise.  I 
am  unaware,  however,  of  any  systematic  monograph  upon 
the  subject  wherein  we  may  find  data  showing  the  char- 
acter of  pathology  in  the  various  animal  groups  or  the 
incidence  of  the  various  lesions.  The  reports  of  some 
zoological  gardens  contain  the  result  of  medical  and 
pathological  data  collected  for  the  report  period.  The 
publication  of  greatest  merit  and  value  is  that  from  the 
Zoological  Society  of  London,  whose  huge  collection 
studied  by  a  large  official  personnel  makes  it  possible  to 
present  valuable  data.  The  New  York  Zoological  Park 
uses  its  material  in  a  similar  manner  and  has  been  able  to 
explain  some  of  the  knotty  problems  so  frequently  met  in 
wild  animal  collections. 

Here  at  Philadelphia  it  has  been  our  practice  now  for 
twenty  years  to  perform  an  autopsy  upon  every  mammal 
and  bird  that  dies,  and  upon  all  of  the  large  or  important 
reptiles.  There  is  no  aquarium  connected  with  the 
Garden.  The  office  of  the  society  keeps  a  record  of  the 
arrival  and  a  general  description  of  every  animal  so  that 
a  brief  history  of  the  specimen  is  usually  available.  The 
keepers  are  required  to  observe  their  charges  regularly 
and  closely  and  to  report  any  abnormalities  to  the  officials 
of  the  Garden.  Somewhat  detailed  discussions  upon  signs 
of  sickness  will  be  given  at  appropriate  places,  where  also 
a  few  remarks  upon  treatment  will  be  included,  but  as  this 
work  is  not  intended  to  be  a  treatise  on  therapy  and  since 
this  subject  does  not  differ  from  that  referable  to  domes- 
ticated animals,  little  space  mil  be  devoted  thereto. 
Upon  death  a  complete  autopsy  is  performed  and  the 


INTRODUCTION  19 

findings  are  recorded  upon  a  printed  form  from  which, 
when  the  histological,  bacteriological  and  parasitological 
studies  have  been  completed,  a  set  of  cross  index  catalogue 
cards  are  typed;  these  are  divided  into  the  principal 
diagnoses  and  determinations.  The  report  of  the  Zoo- 
logical Society,  appearing  at  the  end  of  their  fiscal  year, 
February  28th,  contains  a  resume  of  the  observations  for 
the  year,  together  with  notes  of  interesting  cases  and 
experimental  work. 

There  have  accumulated  the  records  of  nearly  six 
thousand  autopsies  and  upon  them  as  a  basis  has  been 
founded  the  following  report  of  the  incidence  and  nature 
of  pathological  manifestations  in  the  various  animal 
groups,  using  also  as  additional  data,  published  reports 
from  other  gardens.  The  book  might  be  described  as  a 
collection  of  our  studies,  parts  of  which  have  appeared 
as  separate  articles,  but  most  of  which  are  entirely  new, 
put  together  with  as  much  connection  as  the  subject 
matter  will  permit.  The  zoological  and  pathological 
literature  has  been  consulted  very  extensively,  but 
except  for  the  reports  of  zoological  societies  and 
the  publications  of  special  students,  it  usually  rep- 
resents isolated  notes  by  travellers  and  veterinarians 
so  that  many  articles  may  have  been  overlooked. 
Therefore  no  claim  of  perfect  completeness  of  reference 
is  made,  the  statements  resting  chiefly  upon  our  own 
records.  The  subject  will  be  approached  from  the  stand- 
point of  description  and  incidence,  but  it  is  inevitable  that 
comparisons  and  contrasts  must  be  noted. 

Into  the  realm  of  evolution  (1)  I  shall  not  venture 
because  I  appreciate  a  lack  of  adequate  preparation  for 
such  an  attempt,  and  because,  even  if  such  were  not  the 
case,  the  material  at  hand  is  lacking  in  data  upon  fishes, 
many  kinds  of  reptiles  and  invertebrates. 

( 1 )  Those  who  are  interested  in  the  subject  of  disease  in  its  effect  on 
evolution  are  referred  to  Morley  Roberts,  Proceedings,  Zoological  Society, 
London,  1918,  p.  247. 


20     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

A  direct  and  practicable  application  of  these  data  will 
be  in  the  direction  of  explaining  some  of  the  pathological 
states  in  domestic  animals  and  man.  There  are  indeed 
many  disease  entities  or  syndromes  in  these  groups  for 
which  no  useful  hypothesis  has  been  advanced,  while  for 
others  a  partial  explanation  has  been  offered,  usually, 
however,  inadequate  wherewith  to  form  the  basis  of 
rational  prevention  or  therapy.  Thus,  for  example, 
essential  emphysema  seems  to  be  limited  to  the  animals 
of  civilization ;  on  the  other  hand,  the  anatomical  basis  of 
exophthalmic  goitre  may  be  seen  throughout  nearly  the 
whole  animal  kingdom  yet  the  clinical  phenomena  belong 
characteristically  to  man,  and  are  occasionally  seen 
atypically  in  the  dog.  While  it  may  be  impossible  to 
give  a  complete  comparative  anatomy  aiid  physiology  for 
each  of  the  pathological  states,  the  attempt  will  be 
made  to  treat  all  subjects  analogically  through  the  zoo- 
logical orders. 

The  experimental  pathologist  may  find  the  records  of 
the  Garden  useful  in  his  work.  For  example,  he  can  know 
that  rodents  are  not  prominent  among  the  orders  shomng 
spontaneous  arteriosclerosis,  but  that  nephritis  occurs 
among  them  in  about  a  quarter  of  natural  deaths ;  or  he 
may  learn  that  the  Primates  have  a  good  cardiac 
reserve  while  the  Marsupialia  have  not.  Too  often 
experimental  work  is  not  based  upon  facts  including 
natural  probabilities. 

A  collection  of  pathological  data  such  as  is  presented 
in  the  following  pages  may  be  of  assistance  to  vet- 
erinarians and  managers  of  zoological  gardens  in  the 
diagnosis  of  sickness  in  animals,  both  mid  and  domesti- 
cated. We  do  not  presimie  to  offer  a  system  of  veterinary 
medicine,  but  it  is  possible  to  introduce  certain  objective 
findings  of  practical  hygienic  and  therapeutic  value.  Such 
observations  are,  however,  limited  and  in  our  experience 
at  the  Philadelphia  Garden  the  diagnosis  of  disease  in  a 
wdld  animal,  excepting  of  course  those  which  are  per- 


INTRODUCTION  21 

fectly  self-evident,  is  more  often  speculation  and 
conjecture  than  at  all  well  grounded.  It  is  not  uncommon 
for  animals  to  come  to  autopsy  presenting  a  perfect 
galaxy  of  abnormalities,  yet  the  closest  antemortem 
observation  failed  to  reveal  unusual  conduct  or  appear- 
ance. On  the  other  hand  specimens  are  frequently  opened 
whose  organs  fail  to  contain  any  lesions  discoverable 
even  by  careful  study.  Dr.  Henry  Chapman,  sometime 
prosector  to  the  Society,  once  made  a  remark  in  this  con- 
nection— * '  Why  do  they  die  or  how  can  they  live  so  long. ' ' 
Space  is  given  to  this  phase  of  the  observation  of  wild 
animals  in  order  to  emphasize  the  difficulties  of  interpret- 
ing their  conditions,  but  of  course  it  should  be  understood 
that  certain  data  of  value  may  be  gained  by  close  atten- 
tion to  the  details  of  their  normal  behavior  and  to  changes 
which  occur  indicating  that  something  is  wrong. 

The  naturalist  and  the  trained  animal  keeper  are,  in 
our  opinion,  better  judges  of  a  wild  animal's  condition 
than  is  the  veterinarian,  miless  he  be  at  the  same  time  a 
zoologist  and  have  long  experience  with  a  menagerie. 
My  own  observation  of  dogs  and  horses  leads  me  to  think 
that  more  acumen  is  needed  to  interpret  the  actions  of 
wild  animals  since  they  seem  to  have  greater  natural 
reserve,  and  of  course  in  regard  to  them  there  are  many 
more  variables  since  we  see  fewer  specimens  of  each 
species  than  we  know  familiarly  among  domestic  varie- 
ties. The  principal  objects  for  observation  are,  as  in 
veterinary  medicine,  the  eyes,  the  hair  and  skin,  the 
mucous  surfaces,  the  droppings,  the  condition  of  the 
abdomen,  the  appetite  and  the  desire  for  water.  Physical 
examination  is  limited  to  tractable  beasts  and  those  which 
can  be  caught  and  handled  mthout  danger  to  the  per- 
sonnel or  unusual  fright  and  damage  to  themselves.  In 
the  interpretations  of  physical  signs  in  tractable  animals, 
such  as  many  ungulates  and  some  monkeys,  the  experi- 
ence of  the  trained  veterinarian  is  of  the  greatest  value, 
but  this  fails  amongst  carnivores  and  birds.    It  might  be 


22     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

said  that  anesthetics  could  be  used  for  a  thorough  exam- 
ination, but  this  would  be  undesirable  for  a  seriously  sick 
animal  and  it  is,  in  our  experience,  none  too  safe  a  pro- 
cedure although  often  perfectly  practicable.  Animals 
do  not  like  to  be  molested  much  as  they  may  seem  to  enjoy 
attention,  and  when  it  is  possible  it  is  our  practice  to 
avoid  handling  them. 

It  might  be  contended  that  observations  upon  diseased 
states  in  captive  animals  would  not  represent  natural 
developments,  in  other  words,  not  that  which  occurs  in 
the  wild.  Such  indeed  may  be  true  in  regard  to  the  infec- 
tious diseases,  but  since  we  are  imperfectly  informed  as 
to  the  pathology  of  the  wild  state,  we  are  obliged  to  accept 
and  use  the  best  substitute  at  hand.  Moreover  it  seems 
perfectly  fair  to  consider  as  characteristic  for  an  animal 
or  group,  the  physical  and  even  physiological  expressions 
of  morbid  agencies  as  we  know  them,  even  though  the 
animals  be  at  the  time  under  conditions  not  natural  to 
them.  It  would  be  perhaps  incorrect  to  say  that  cirrhosis 
of  the  liver  occurs  in  .6  per  cent,  of  animals  in  the  wild 
as  is  the  case  for  our  autopsies,  since  incorrect  food  and 
infections  are  potent  in  its  causation ;  on  the  other  hand, 
our  experience  and  some  few  data  from  naturalists  and 
pathologists  make  it  conceivable  that  tumors  occur  to  this 
number  in  native  states.  The  incidence  of  tumors  in  mid 
rodents  is  quite  well  known.  Degenerations  and  fibroses, 
the  result  of  parasitism,  are  known  to  exist  throughout 
the  entire  animal  kingdom.  Further  to  illustrate  how 
pathology  is  distributed  in  wild  life,  Plimmer's 
experience  (2)  with  500  rats  {M.  decumanus)  might  be 
cited.  He  found  the  following:  Tubercle  3  times,  tape 
worm  cysts  10,  Tryp.  lewisi  49,  empyema  2,  tumor  of  jaw 
from  old  injury  1,  pleuritis  and  hydrothorax  1.  Bacteria 
were  found  in  71  rats,  40  times  in  the  lungs,  31  times  in 
the  spleen;  saccharomyces  were  found  16  times  in  the 
lungs.    Dr.  W.  L.  Abbott  reports  to  us  personally  that  he 

(2)   Proceedings,  Zoological  Society,  London,  1911. 


INTRODUCTION  23 

has  repeatedly  found  coiled  exproventricular  worms  in 
the  wild  specimens  he  has  collected.  Not  only  are  we 
informed  of  some  isolated  and  individual  pathological 
states  but  the  existence  of  epizootics  of  communicable 
disease  among  wild  life  is  well  authenticated.  The 
simple  citation  of  the  extermination  of  deer  in  one  section 
of  Colorado  by  pleuropneumonia  will  suffice  to  illustrate 
this  point.  Other  examples  are,  however,  interesting. 
The  occurrence  of  changes  in  the  jaw  bone  ahnost 
certainly  those  of  actinomycosis  is  reported  by 
Blair,  the  specimens  being  shot  in  the  mid  and  believed 
never  to  have  been  near  civilization.  The  white-tailed 
deer  of  the  Swan  River  Valley  in  Montana,  are  knomi  to 
be  constant  carriers  of  liver-flukes. 

It  would  seem  therefore  that  it  is  not  unfair  to  use 
material  gathered  from  animals  under  somewhat 
unnatural  conditions  as  representing  the  reaction  of  the 
zoological  orders  to  pathogenic  agencies.  Such  conclu- 
sions must  however  be  made  very  guardedly,  for 
it  is  probable  that  not  over  ten  per  cent,  of  the 
total  number  of  mammalian  and  avian  species  are  to 
be  observed  in  captivity.  Because  of  the  number 
of  orders  and  the  great  variety  of  genera  included  in 
the  present  study  it  is  probable  nevertheless  that 
the  lesions  are  fairly  representative  of  the  whole 
animal  kingdom. 

However,  the  nimibers  and  percentages  given  should 
be  read  to  indicate  the  probabilities  and  should  not 
be  interpreted  as  implying  the  mortality  relationships 
smce  different  varieties  have  differing  powers  of 
resistance  to  the  same  pathological  state.  The  margin 
of  safety  in  any  given  group  for  one  or  several  dif- 
ferent disease  entities  cannot  at  present  be  stated  mth 
any  degree  of  precision  but  this  factor  is  doubtless  very 
great.  The  work  of  physiologists  suggests  that  there  is  a 
reserve  power  in  the  human  lung  sufficient  to  sustain  life 
until  five-sixths  of  the  functionating  organ  is  useless,  and 


24     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

I  shall  quote  a  case  of  an  opossum  wherein  only  one-tenth 
of  the  respirable  surface  seemed  to  have  remained;  we 
have  repeatedly  seen  both  lungs  of  a  monkey  apparently 
entirely  solid.  Such  physical  vital  incompatibilities 
might  be  exemplified  by  many  other  cases,  but  when  one 
reviews  the  physiological  margin  of  safety,  inexplicable 
and  contradictory  instances  are  equally  numerous.  1 
have  seen  a  male  deer  run  a  doe  against  the  fence  and 
butt  her,  without  result,  whereas  in  an  apparently  similar 
occurrence  the  animal  would  be  dead  in  a  short  time. 
Numerous  instances  of  slight  enteritis  of  a  short  stretch 
of  duodenum  or  ileum  have  killed,  with  almost  nothing  to 
be  found  microscopically,  and  on  many  occasions  we  have 
been  chagrined  in  being  unable  to  discover  the  cause  of 
death.  The  capacity  of  self-healing  is  a  variable  one,  but 
seems  in  direct  proportion  to  the  quietness  and  seclusion 
possible  for  the  animal  and  inversely  to  the  chance  of 
bacterial  infection. 

The  effect  of  captivity  has  been  the  subject  of  much 
speculation.  For  the  preservation  of  health  it  would  seem 
that  animals  require  periods  of  rest  and  activity,  thor- 
ough elimination,  possibly  a  moderate  exercise  of  their 
procreative  functions,  but  most  of  all,  appropriate  food 
obtained  by  the  physical  effort  we  term  chase.  All  but 
the  very  last  condition  is  supplied  in  a  measure  in  well 
managed  collections.  The  degenerating  effect  of  the 
absence  of  chase  must  be  admitted.  An  interesting  and 
suggestive  example  of  this  was  noted  by  Mr.  Jones  at  the 
London  Zoological  Gardens.  He  observed  the  skull  of  a 
lion  that  had  been  in  captivity  thirteen  years,  in  which  the 
canine  area  of  the  face  and  the  part  of  the  skull  acting  as 
the  insertion  for  the  seizing  and  holding  muscles  had 
undergone  atrophy  while  the  chewing  muscles  with  their 
bony  bases  had  remained  normal.  Numerous  examples 
of  disease  atrophy  are  on  record  and  those  of  a 
physical  nature  must  have  counterparts  in  the  realm 
of  physiology.    The  size  to  which  an  animal  will  attain 


INTRODUCTION  25 

cannot  be  estimated  by  the  examples  seen  in  menageries. 
Judging  by  the  accounts  of  collectors  and  hunters  and 
upon  the  more  reliable  of  the  moving  picture  displays  of 
wild  animals  in  their  native  haunts,  it  would  seem  prob- 
able that  under  normal  conditions  of  habitat  the  average 
size  of  wild  beasts  is  considerably  in  excess  of  that  in 
park  specimens. 

The  effect  of  captivity  may  also  be  felt  in  the  direction 
of  reduced  resistance  to  infectious  diseases.  Brooks,  of 
the  New  York  Park,  expresses  the  view  that  captivity 
increases  susceptibility  to  bacteria  and  causes  paren- 
chymatous degenerations.  In  the  latter  direction  it  is 
interesting  to  learn  that  Seligman  of  London  claims  to 
have  seen  sudden  deaths  in  wading  and  struthious  birds 
from  myocardial  disease,  without  valvular  or  other 
lesions,  for  which  he  holds  the  enervating  effects  of  cap- 
tivity responsible.  It  is  well  recognized  that  a  species 
may  be  unusually  susceptible  to  a  disease  that  it  has  not 
encountered  in  its  phylogenic  development.  Man  illus- 
trates this  peculiarity  very  clearly.  Europeans  were 
found  exceedingly  susceptible  to  sleeping  sickness  when 
they  went  first  to  the  part  of  Africa  inhabited  by  the  tsetse 
fly,  and  the  American  Indians  died  in  hordes  when  they 
met  the  tubercle  bacillus  for  the  first  time.  Judging  by 
the  ravages  of  tuberculosis  in  captive  monkeys  a  similar 
susceptibility  probably  explains  the  matter  for  there  are 
no  entirely  satisfactory  records  of  this  disease  among 
them  in  the  wild  state. 

In  so  far  as  general  susceptibility  to  infection  is  con- 
cerned, it  may  be  in  part  due  to  one  of  the  artificial 
conditions  of  captivity,  that  of  inbreeding.  This  influence 
is  undoubtedly  very  great,  both  by  chance  in  families,  and 
by  intention  on  the  part  of  dealers  as  well  as  the  mating 
which  occurs  in  menageries.  However,  it  is  not  known 
how  far  inbreeding  may  go  in  the  wild  state  so  that  one 
must  be  very  careful  about  drawing  conclusions  in  this 
particular.    Several  years  ago,  at  the  time  we  reported 

3 


26     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

the  neoplasms  found  at  the  Garden,  discussion  arose  as 
to  the  effect  of  inbreeding,  and  thereafter  some  observa- 
tions were  made  in  this  direction.  With  the  exception  of 
the  hyperplasias  of  the  thyroid,  not  certainly  of  neo- 
plastic nature,  in  a  much  mixed-up  family  of  wolves,  we 
could  find  no  evidence  that  inbreeding  was  responsible 
for  tumors.  Plimmer  and  Murray  of  London,  seem  to 
imply  that  some  of  their  inbred  animals  are  likely  to  have 
tumors ;  reference  to  this  matter  will  be  made  later  in  this 
book.  In  so  far  as  diseases  of  the  organic  systems  are 
concerning  those  of  the  bones  seem  to  be  the  only  ones  in 
which  inbreeding  is  significant. 

The  individual  resistance  will  be  reduced  of  course  by 
the  unsanitary  surroundings  incident  to  trapping,  ship- 
ping and  storage,  but  this  need  not  affect  the  figures  or 
pathological  tendencies  of  classes  or  orders. 

The  effect  of  captivity  is  felt  in  another  way.  A  very 
large  percentage  of  wild  life  perishes  during  the  first 
weeks  or  months  after  its  capture,  and  in  gardens  the 
heaviest  mortality  occurs  among  the  recent  arrivals.  The 
London  Garden  figures  that  from  thirty-three  per  cent,  to 
fifty  per  cent,  of  their  total  mortality  is  in  animals  that 
have  not  been  in  the  garden  six  months  and  that  die 
because  they  are  not  yet  accustomed  to  their  new  sur- 
roundings. It  seems  to  us,  both  from  an  academic 
and  a  practical  standpoint,  that  this  is  a  long 
time  and  should  afford  ample  opportunity  for  the 
garden  to  study  the  specimen  and  for  the  specimen  to 
become  acclimated.  These  early  deaths  are  perhaps 
to  be  ascribed  in  large  part  to  failure  of  acclimatiza- 
tion but  many  are  doubtless  the  result  of  infection 
acquired  in  the  wild,  in  transit  while  in  the  hold  of  ves- 
sels, at  quarantine,  or  in  trains,  or  at  the  establishments 
of  dealers.  We  have  seen  a  few  deaths  which  have  fol- 
lowed behavior  that  might  be  likened  to  homesickness. 
Perhaps  the  age  at  arrival  has  an  influence  upon  the 
morbidity  and  mortality  of  wild  animals,  for  it  is  easily 


INTRODUCTION  27 

conceivable  that  the  young  and  the  very  old  might  adapt 
themselves  to  new  surroundings  with  much  less  readiness 
than  the  sturdy  middle-aged  adult.  The  age  of  animals 
upon  arrival  is  verj^  rarely  knowm,  and  can  only  be 
recorded  as  "young,"  "fully  developed,"  and  "old." 
This  will  have  an  effect  upon  statistics  and  when  possible 
is  noted  in  the  text,  but  this  is  not  practicable  to  the 
extent  we  desire.  The  meaning  of  "young,"  "adult," 
and  "old"  is  not  the  same  throughout  the  animal  orders 
nor  even  \vithin  orders. 

Mitchell  (3)  has  attempted  to  gain  concrete  ideas 
of  the  expectancy  of  life  among  animals  by  analyz- 
ing the  records  of  the  London  Gardens.  This  gentleman 
bases  his  figures  upon  kno^vn  ages  and  the  length 
of  time  in  captivity,  from  a  combination  of  which 
data  the  specific  viability  and  the  potential  longevity 
may  be  estimated.  Such  results,  he  admits,  can 
only  be  approximate  and  they  show  within  classes  and 
orders,  a  decided  lack  of  uniformity.  The  terms  "spe- 
cific" and  "potential"  longevity,  coined  by  Sir  Ray 
Lankester,  apply,  for  the  first,  to  the  average  length  of 
life  as  it  is  affected  by  external  conditions  and  those 
incident  to  procreation,  while,  if  an  animal  be  under  ideal 
conditions  it  will  attain  the  potential  longevity  which  is 
longer  than  the  former.  These  considerations  have  a 
biological  and  economic  importance,  while  a  knowledge 
of  the  pathology  shown  by  the  various  groups  may  help 
to  explain  these  durations  of  life.  Contrariwise  figures 
of  the  expected  longevity  may  assist  us  in  evaluating 
youth  and  senility  in  the  causes  of  death  but  can  hardly 
affect  the  comparative  nature  of  the  lesion. 

A  resume  of  Mitchell's  studies  indicates  that  the 
higher  apes  have  a  potential  longevity  and  a  hardihood 
much  less  than  man  but  still  upwards  of  thirty  years.  As 
one  investigates  lower  in  monkeys,  life  periods  become 
shorter,  while  in  the  next  order,  Lemures,  the  length  of 

(3)   Proceedings,  Zoological  Society,  London,  1911,  p.  425. 


28     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

life  rises.  Carnivora  have  a  reasonably  good  vitality, 
their  potential  periods  varying  from  ten  years  in  the 
foxes  to  thirty-three  years  in  bears.  Insect  eating 
animals  are  short  lived,  three  years  being  a  maximum. 
The  Bat  family  shows  great  variations,  the  greatest  life 
being  not  over  seventeen  years.  The  Rodentia  have  long 
lives  compared  to  their  sizes — twenty  years  in  porcupines, 
fifteen  years  in  squirrels,  thirteen  in  marmots,  nine  in 
agoutis  and  capybaras,  and  three  in  dormice  (which  is 
also  about  the  maximum  for  the  rat).  Hy races  live  four 
years  on  the  average.  Proboscidea,  although  reputed  to 
live  to  great  age,  probably  rarely  live  a  half  century  and 
may  be  said  to  have  an  expectancy  of  twenty  to  thirty 
years.  Perissodactyla  (horses,  tapirs  and  rhinoceroses) 
while  they  may  live  half  a  century,  have  an  average  life 
of  between  fifteen  and  thirty  years.  The  closely  related 
Artiodactyla  fall  into  two  groups,  a  first  comprising 
antelopes,  sheep,  goats  and  deer  which  rarely  exceed 
seventeen  years,  and  a  second  consisting  of  cattle,  camels 
and  giraffes,  which  vary  in  expectancy  from  eighteen  to 
thirty  years.  The  smaller  members  of  the  Ungulata  have 
in  relation  to  size  a  relatively  greater  viability,  the 
ruminants,  however,  having  on  the  whole  a  low  viability. 
Marsupials  vary  from  a  maximum  of  seven  years  in  the 
opossum  to  eighteen  in  the  wombats,  but  none  of  this 
group  has  a  good  viability.  The  Aves  as  a  class  or  if 
compared  according  to  dietary  requirements,  have  longer 
potential  ages  and  better  viability  than  mammals.  Pas- 
serine birds  average  twenty  years  and  many  live  to  sixty, 
while  the  Picariae  approach  the  former  figure  but  do  not 
have  such  good  viability.  Psittaci  and  Striges  may  live  a 
half  century  but  the  resistance  of  the  latter  is  much 
reduced  by  any  unfavorable  surroundings.  The  rapta- 
tory  birds  live  fifty  years,  but  their  viability  is  variable. 
Herodiones  have  a  maximum  expectancy  of  thirty  years 
and  good  resistance,  while  their  relatives,  Steganopodes, 
may  live  fifty  years,   and  Odontoglossae  have   a   good 


INTRODUCTION  29 

viability,  up  to  twenty  years.  Anserine  birds  may  live  to 
be  fifty,  and,  unless  conditions  are  quite  unfavorable, 
have  a  good  resistance.  Columbae  may  under  good  con- 
ditions live  to  be  fifty.  Gallinaceous  birds  may  only  be 
expected  to  survive  twenty  years,  a  figure  also  given  for 
Fulicariae.  Alectorides  may  live  up  to  fifty  years. 
Limicolas,  though  they  do  not  thrive  in  captivity,  may  live 
thirty  years.  Impennes  live  poorly  under  artificial  con- 
ditions, the  greatest  record  being  twelve  years,  a  figure 
also  holding  for  Crypturi.  Struthiones,  if  the  conditions 
be  right,  may  live  fifty  years. 

Because  of  the  variable  specific  longevities,  it  is  fre- 
quently difi&cult  to  decide  when  an  animal  is  senile.  Man 
is  said  to  be  as  old  as  his  arteries,  and  his  span  of  life 
nowadays  is  in  the  neighborhood  of  half  a  century.  Par- 
rots exhibit  lesions  of  the  vascular  system  comparable 
to  the  arteriocapillary  fibrosis  of  human  beings,  and  their 
expected  longevity  is  about  the  same  or  a  little  greater. 
From  a  study  of  our  cases  of  this  lesion  in  parrots  it  can 
be  said  to  appear  quite  early  in  life  and  not  to  lead  to 
organic  disease  as  it  is  alleged  to  do  in  man.  It  is,  how- 
ever, interesting  to  note  that  in  those  animals  which  are 
supposed  to  have  the  longest  specific  lives — elephants, 
snakes,  anserine  and  raptatory  birds,  parrots — there  is 
relatively  low  mortality  and  fewer  infectious  diseases 
are  encountered.  The  last  part  of  this  statement  should 
be  qualified  by  stating  that  anserine  birds  and  parrots 
are  quite  susceptible  to  mycoses,  in  all  probability  from 
musty  food,  which  raises  their  death  rate,  but  as 
this  is  accidental  and  artificial,  it  can  be  excluded 
from,  consideration. 

In  a  rough  way  there  is  a  direct  relationship  between 
the  size  of  an  animal  and  its  longevity,  but  this  is  not  close 
enough  to  be  a  reliable  guide ;  whales  and  elephants  live  a 
long  time,  but  so  do  snakes  and  parrots.  Within  orders 
this  relation  of  size  and  expected  longevity  is  more  easily 
seen  but  is  not  absolute.  I  cannot  state,  according  to  my 


30     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

present  studies,  that  there  is  an  unqualified  relationship 
between  the  size  and  expected  longevity  of  an  animal  and 
its  pathological  lesions. 

The  immediate  surroundings  and  the  management  of 
captive  animals  have  a  very  direct  and  important  bearing 
upon  the  mortality  and  perhaps  upon  the  incidence  of 
morbid  processes  but  probably  not  upon  the  character  of 
the  latter.  A  full  knowledge  on  the  part  of  the  personnel  of 
a  zoological  society  concerning  the  habits  and  habitat  of 
eveiy  animal  in  their  keeping  is  essential,  to  wliich  must 
be  added  a  group  of  interested  keepers.  In  engaging  the 
last,  it  should  not  be  forgotten  that  certain  men 
have  ''a  way"  with  animals  and  that  others  cannot 
manage  themselves. 

The  enormous  literature  at  the  disposal  of  the 
naturalists  permits  executive  officers  to  formulate  a  plan 
of  housing  and  feeding  with  fair  accuracy  for  each  kind 
of  animal,  but  of  course  it  is  rarely  possible  to  obtain  in 
sufficient  quantity  the  natural  food  {e.g.,  ants  for  ant- 
eaters).  In  so  far  as  food  is  concerned  it  seems  that  with 
a  few  exceptions  like  the  one  just  mentioned,  the  substitu- 
tions made  at  the  zoological  gardens  are  nearly 
satisfactory.  The  elements  in  which  the  captive  diet  is 
poor  are  the  inorganic  salts  and  vitamins  since  Dr. 
Corson- A\Tiite,  some  of  whose  work  is  included  in  a  later 
chapter,  has  shown  that  for  those  animals  which  our 
statistics  indicate  as  most  prone  to  have  rickets  and  osteo- 
malacia, the  available  phosphorus  and  calcium  are  low, 
and  one  vitamin  was  also  below  the  desired  quantity.  In 
this  regard,  however,  I  am  not  at  all  con\inced  that  diet 
alone  mil  suffice  to  explain  these  degenerative  osseous 
diseases ;  I  shall  take  this  up  more  fully  later.  Careful 
inspection  of  all  food  should  be  made  and  cleanliness 
(sifting  of  cereals,  protection  of  meat  from  flies,  etc.),  is 
indispensable.  The  mortality  among  our  camivora  has 
materially  decreased  since  the  horse  meat  after  butcher- 
ing was  placed  in  covered  galvanized  iron  pans.    There 


INTRODUCTION  31 

are  many  problems  of  feeding,  too  numerous  to  be  cov- 
ered in  a  survey  of  this  sort,  which  must  be  solved,  and 
it  is  a  credit  to  superintendents  that  this  they  have 
studied  carefully. 

There  are  two  problems  in  the  management  of 
animals  upon  which  much  difference  of  opinion  exists, 
namely  the  heating  of  houses  and  the  material  of  which 
cages  are  made.  It  seems  to  be  the  practice  in  many 
gardens  to  keep  animals  very  warm.  Dr.  Chalmers 
Mitchell  states  unqualifiedly  that  adult  animals  do 
not  have  to  be  kept  warm,  and  that  even  an  equable 
temperature  is  not  demanded,  variations  in  temperature 
having  a  distinctly  stimulating  effect.  However  he  main- 
tains that  they  should  be  kept  dry  and  must  be  supplied 
with  a  shelter.  This  is  in  accord  with  the  experience  at 
the  Philadelphia  Garden,  since  for  many  years  we  have 
allowed  access  to  the  open  air  all  winter  to  every  animal 
that  could  stand  it.  A  large  group  of  macaques  has  now 
lived  entirely  in  an  open  ''band  stand"  cage  for  nine 
years  with  a  lower  mortality  than  in  the  rest  of  the 
monkey  collection,  which  is  permitted  to  go  indoors  some 
of  the  time.  Occasionally  one  in  poor  health  is  frozen  to 
death,  and  healthy  ones  may  lose  fingers,  toes,  or  a  part 
of  the  tail,  but  the  general  condition  is  so  much  improved 
that  they  present  an  attractive  exhibit  to  visitors.  Unless 
a  storm  be  of  great  severity,  wild  animals  are  usually 
indifferent  to  it  although  they  may  seek  their  shelter. 
Snow  apparently  is  no  source  of  fear  to  them,  and  many 
enjoy  playing  in  it.  The  general  principles  of  the  enclosure 
should  be  proper  lighting,  free  access  of  air,  dryness  and 
shelter  in  time  of  storm,  the  last  so  arranged  that  the 
sleeping  place  is  well  protected.  Appropriate  arrange- 
ments should  be  made  for  nocturnal  animals,  regardless 
of  their  visibility  to  visitors,  if  their  preservation  is 
of  importance. 

The  hygiene  of  communicable  disease  has  influenced 
everyone  to  use  concrete  and  metal  for  cages.  These  sub- 


32     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

stances  are  without  doubt  most  simply  kept  clean,  but 
they  are  heat-conducting  and  remain  cold  or  damp  longer 
than  wood  or  the  ground.  It  may  be  claimed  that  the 
latter  two  cannot  be  disinfected  so  well,  but  this  need  not 
militate  against  their  use.  Wood  can  be  disinfected  by 
sunlight  or  by  mechanical  cleaning  plus  disinfectants,  by 
a  blast  lamp  and  by  paint.  The  ground  will  disinfect 
itself  if  allowed  to  lie  fallow  for  a  time,  or  it  may  be 
turned  over  after  sprinkling  with  lime.  It  is  fair  to  note 
that  the  New  York  Zoological  Garden  reduced  their  mor- 
tality, especially  from  verminous  pneumonia,  by  changing 
some  deer  herds  to  concrete  paved  enclosures;  if  that 
were  the  only  change  made  the  result  would  be  very 
significant,  but  it  should  not  be  forgotten  that  another 
clean  ground  range  might  have  served  as  well  to  a  herd 
from  which  the  infected  ones  had  died.  My  own  observa- 
tions with  guinea-pigs,  rabbits,  mice  and  dogs  lead  me  to 
believe  that  they  thrive  and  breed  better  on  wooden  floors 
than  on  metal  or  stone. 

I  have  tried  to  work  out  figures  to  show  that  more 
animals  die  when  housed  in  enclosures  of  stone  and  metal 
than  when  upon  the  earth  or  on  wood,  but  the  attempt  has 
been  unsuccessful  chiefly  because  of  the  presence  of  epi- 
demics and  parasites,  principally  among  the  birds.  The 
attempt  was  further  embarrassed  because  some  members 
of  an  order  are  housed  on  both  floorings.  However,  there 
was  no  great  advantage  for  the  metal  and  concrete  floors 
even  after  the  epidemic  had  been  discounted.  This  Gar- 
den does  not  have  a  great  number  of  pneumonias,  a 
disease  said  to  be  favored  by  dampness  and  cold,  but 
those  that  occur  are  chiefly  among  the  small  mammals,  on 
wooden  floors  and  in  the  large  bird  house  in  cages  of  con- 
crete and  metal.  However,  the  construction  of  both  these 
houses  permits  the  visitors  to  approach  very  close  to  the 
cage,  a  factor  that  doubtless  explains  the  dispropor- 
tionate incidence  of  inflammation  of  the  lungs.  In  so  far 
as  outdoor  fowl  and  ungulate  ranges  are  concerned,  they 


INTRODUCTION  33 

should  be  changed  frequently  under  the  best  conditions 
since  occasionally  one  will  find  groups  doing  badly  until 
moved.  Moreover  the  ground  becomes  contaminated 
with  parasites  such  as  esophagostomum  and  heterakis, 
infestation  with  which  while  not  very  serious  in  itself, 
may  lead  to  fatal  infection  with  bacteria. 

The  effect  of  animal  parasites  upon  the  morbidity  and 
mortality  of  wild  beasts  and  birds  in  captivity  is  by  no 
means  clear,  and  Doctor  Weidman  aiid  I  are  inclined  to 
be  sceptical,  with  certain  reservations  of  course,  of  their 
great  importance  in  the  death  rate.  Doctor  Weidman  has 
kindly  agreed  to  contribute  a  chapter  upon  the  general 
distribution  of  protozoal  and  metazoal  parasites  with  a 
summary  of  their  probable  pathogenic  importance. 

The  groups  known  to  have  a  decided  pathological 
power  might  be  divided  into  the  toxic,  the  tumor  formers 
and  the  mechanically  obstructive ;  certain  parasites  have 
properties  placing  them  in  two  of  these  classes.  The  first 
group  comprises  the  hemosporidia  and  hemogregarines, 
the  uncinaria  and  some  of  the  cestodes,  forms  which  pro- 
duce hemolysis  and  hemorrhages  with  varying  grades  of 
anemia.  The  importance  of  this  group  is  shown  chiefly 
among  the  Aves,  in  which  high  grades  of  anemia  are 
occasionally  met  from  malarial  infections,  but  cats  and 
dogs  or  even  herbivores  also  frequently  suffer  from 
hookworm.  The  tumor-producers  are  chiefly  echinococ- 
cus  worms,  the  cysts  of  which  may  grow  large  enough  to 
occupy  nearly  the  entire  abdomen.  A  certain  grade  of 
anemia  and  general  ill  health  accompany  this  hydatid 
disease,  partly  the  result  of  a  toxin  and  partly  by  damage 
to  important  viscera.  Those  parasites  which  obstruct 
mechanically  do  so  by  their  own  bulk  or  by  an  accom- 
panying inflammation,  incited  by  them  as  foreign  bodies 
or  by  bacteria  which  have  gained  entrance  at  the  irritated 
point.  This  is  exemplified  by  the  enormous  collection  of 
nematodes  sometimes  found  in  reptiles  (a  pailful  was 
removed  from  a  python)   and  by  the  tightly  coiled  or 


34     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

tangled  thread  and  tape  worms  frequently  found  in  birds. 
The  effect  of  swelling  by  the  mucous  membrane  under  the 
influence  of  worms  is  illustrated  by  the  infestation  of  the 
proventricle  in  parrots.  Here  spiroptera  penetrate  into 
and  under  the  glandular  layer  which  sweUs  and  pours  out 
mucus,  the  total  mass  of  nematodes,  mucus  and  tissue 
obstructing  the  passage. 

Very  many  animals  show  parasitic  infestation  at  post- 
mortem, but  the  percentage  in  which  they  can  be  said  to 
be  principal  causes  of  death  is  quite  small,  while  that  in 
which  they  play  a  role  as  activator  of  the  terminal  condi- 
tion is  also  small  but  indeterminate.  The  latter  group 
comprise,  together  with  the  anemias  mentioned  above, 
certain  forms  of  pneumonia,  of  hepatic  and  vascular 
lesions.  Inflammations  of  the  lungs  from  ascaris  and 
paragonimus  are  fairly  well  knowai ;  fortunately  we  have 
been  troubled  less  with  this  than  have  most  gardens,  pos- 
sibly because  we  do  not  have  such  large  herds  of  herbivora 
susceptible  to  it.  Hepatic  diseases  from  flukes,  from  coc- 
cidia  and  from  amoebae  we  have  always  with  us  in  small 
numbers,  but  they  are  unimportant  excepting  enterohep- 
atitis,  a  condition  which  appears  in  nearly  all  orders. 
This  last  disease,  be  it  purely  amoebic  as  in  dysentery  of 
man  and  monkeys,  or  like  blackhead  of  turkeys  and  chick- 
ens or  in  the  forms  of  quail  disease,  arrests  the  attention 
at  once  and  evokes  a  desire  to  explain  the  association  of 
large  intestines  and  liver.  Parasitic  vascular  lesions  are 
relatively  unimportant. 

Taking  parasitic  infestations  by  and  large,  there  are 
close  similarities  throughout  the  entire  animal  kingdom, 
and  the  effects  produced  by  a  given  genus  will  be  repeated 
almost  exactly  in  several  others.  The  pathological  pic- 
tures of  anemia,  of  hepatic  degeneration,  of  cystic 
degeneration,  of  colonic  ulceration  or  of  fibroses  are  simi- 
lar in  different  hosts,  only  slight  variations  in  the  type  of 
inflammation  being  noted,  for  instance  in  reptiles  and 
birds  as  against  the  mammals.  We  have  made  rather  close 


INTRODUCTION  35 

observations  upon  the  effect  of  parasites  in  the  produc- 
tion of  neoplasms,  incited  by  Fibiger's  discovery  of 
nematodes  in  the  rat 's  stomach  cancer,  but,  with  the  pos- 
sible exception  of  a  papillomatous  growth  in  the  stomach 
of  opossums  from  the  action  (?)  of  physaloptera,  we  have 
been  unable  to  establish  such  an  etiological  relationship. 
A  decision  of  the  importance  of  parasites  in  any  given 
case  is  not  without  its  difficulty,  and  we  are  inclined  to 
reserve  judgment  pending  further  analysis  unless  the 
effect  of  the  invaders  is  unequivocal.  Leiper  (4) 
does  not  seem  to  credit  animal  parasites  with  a 
great  effect  on  the  mortality  after  a  specimen  has 
been  in  the  collection  six  months  since  all  the 
intestinal  varieties  he  studied  came  from  animals  dying 
in  that  period.  On  the  other  hand  the  forms  which 
invaded  the  internal  organs  and  tissues  were,  in  his 
series,  from  specimens  resident  several  years  in  the 
garden.  He  seems  to  think  the  conditions  of  life  at  the 
garden  favor  the  expulsion  of  intestinal  worms.  To  what 
extent  some  intestinal  worms  may  be  commensal  remains 
as  uncertain  as  the  value  of  certain  bacteria  in  the  gut 
tract.  In  man  considerable  importance  has  been  ascribed 
to  certain  fermentative  and  putrefactive  germs  in  the 
maintenance  of  a  reaction  unfavorable  to  strict  pathogens 
and  some  observers  have  looked  at  them  as  possessing  a 
digestive  power.  In  the  digestive  tract  of  the  animals 
eating  large  quantities  of  carbohydrate  as  cellulose, 
nature  provides  for  its  use  by  rumination  and  by 
supplying  a  large  hind  gut,  by  which  means  secondary 
mastication  and  bacterial  decomposition  of  the  cellulose 
capsule  insures  its  fuU  use.  Possibly  a  similar  usefuhiess 
may  be  finally  ascribed  to  some  animal  microbes  or  even 
larger  metozoa. 

The  role  of  vegetable  parasites  in  the  causation  of  dis- 
ease among  wild  animals  seems  as  undoubted  as  it  is  in 
the  human  being  and  the  pathologic  results  are  usually  as 

(4)   Proceedings,  Zoological  Society,  London.    1911,  p.   620. 


36     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

clear,  at  least  for  the  entities  of  which  we  have  exact  data, 
based  upon  comparisons  with  man  and  domestic  animals. 

There  seems  to  be  no  essential  difference  among 
mammals  between  the  pathological  pictures  of  infectious 
septicemias,  the  mucous  and  serous  membrane  inflam- 
mations and  tuberculosis  for  example.  They  are 
characterized  by  fibrinous,  purulent  or  infiltrative 
inflammations  which  may  go  on  to  necrosis  or  repair,  by 
fever,  by  leucocytosis  and  by  evidences  of  resistance — all 
of  these  things  occurring  in  a  similar  way  throughout  the 
class.  Of  course  not  all  animals  are  receptive  to  all 
infections  since  specific  racial  and  generic  immunities 
exist,  but  the  basic  response  in  terms  of  pathology  is 
similar.  There  are  no  normal  means  of  judging  the  sus- 
ceptibility of  wild  animals  on  their  native  heath  to  the 
important  pathogens  of  civilization,  pneumococci,  strep- 
tococci, staphylococci,  cholera  bacilli,  the  typhocolon 
group,  the  Friedlander  group  and  others,  but  it  is  inter- 
esting to  note  that  in  captive  conditions  they  evince  some 
receptivity  to  these  germs  or  their  congeners.  The  pneu- 
mococcus  takes  a  fairly  heavy  toll  in  zoological  collections 
every  year  and  the  Friedlander  bacillus,  not  a  very  com- 
mon cause  of  human  pneumonitis,  has  been  seen  here  and 
at  London. 

Among  the  birds,  however,  quite  distinct  differences 
in  some  pathological  processes  occur,  not  only  from  the 
mammals  but  also  within  the  class.  '  As  a  whole  birds 
do  not  produce  pus  as  we  know  it  in  man,  probably 
because  of  the  absence  from  their  leucocytes  of  a 
protein-splitting  ferment;  their  leucoc>i;e-producing 
organs  do  not  seem  to  respond  as  readily  to  a  virus, 
the  place  of  purulent  exudate  being  taken  by  a  coagu- 
lum  or  necrosis.  The  former  varies  from  a  clear 
gelatin-like  material  seen  upon  serous  surfaces  to  a  thick 
mat  or  mass  of  coarse  but  short  fibrinous  strands. 
Necrosis  may  succeed  upon  the  latter  or  occur  so 
promptly    as    to    appear    like    the    original    form    of 


INTRODUCTION  37 

degeneration.  It  is  usually  rapid,  accompanied  by  a 
circumferential  congestion  but  not  associated  with  active 
phagocytosis.  Giant  ceU  production  is  variable,  but  when 
developed  the  appearance  is  like  that  of  large  syncytia. 
Hemolysis  is  not  marked  in  the  simple  infections  but  a 
hyperplasia  of  the  mononuclear  nodes  of  the  liver  is  the 
rule.  The  function  of  this  nodal  increase  is  not  quite 
clear.  It  has  been  always  thought  that  the  scanty  bone 
marrow  would  supply  the  necessary  erythrocytes,  but  we 
have  seen  these  mononuclear  areas  full  of  pale  red  cells 
fitted  with  round  nuclei  and  without  pigment.  The  fibrin 
mentioned  above  does  not  have  the  delicate  interweaving 
that  we  know  in  a  fibrinous  exudate  in  man.  This  is  inter- 
esting when  we  consider  the  composition  of  the  blood 
and  its  coagulation  in  the  Aves.  The  cell  upon  which 
human  coagulation  seems  to  depend,  the  platelet,  is  rep- 
resented in  birds  by  the  thrombocyte,  which  appears  only 
up  to  about  50,000  per  cubic  millimetre.  Coagulation  time 
is  relatively  short  and  the  resulting  clot  is  firm  and 
irregular.  Perhaps  this  may  have  something  to  do  with 
the  nature  of  an  inflammatory  exudate. 

The  response  to  infection  on  the  part  of  birds  may  to 
some  extent  depend  upon  differences  in  anatomy,  which 
are  quite  distinct,  not  only  from  the  mammals  within 
which  class  the  anatomy  is  more  uniform,  but  also  from 
one  avian  order  to  another.  These  differences  among  the 
birds  may  be  exemplified  by  the  large  foramina  between 
lungs  and  air  sacs  in  the  water  birds,  a  passage  which 
permits  infection,  notably  mycosis,  to  spread  from 
the  first  to  the  second.  Again  the  close  apposition  of 
the  pancreas  to  the  duodenum  over  a  long  stretch 
permits  easy  infection  of  the  former  from  the  latter. 
Still  again  the  large  renal-portal  vein  in  the  gallinaceous 
birds  explains  some  of  the  infections  of  the  liver  sec- 
ondary to  intestinal  disease.  The  position  of  the  lungs, 
deep  in  the  thorax  and  fitted  into  recesses  made  by  the 
sharp  anterior  border  of  the  ribs  and  overlaid  anteriorly 


38     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

by  a  rather  firm  air  sac  wall,  makes  it  difficult  for  these 
organs  to  expand  and  therefore  renders  even  a  simple 
congestion  a  dangerous  thing.  The  position  of  the  ovary 
subjects  the  shell-less  egg  to  much  danger  from  the 
intestinal  area. 

These  and  many  other  peculiarities  of  anatomy 
affect  the  pathological  picture  in  birds.  To  be  sure 
there  are  also  noteworthy  differences  among  the  Mam- 
malia, notably  in  the  intestinal  and  genital  tracts, 
but  the  pathologic  response  is  not  so  varied  as  in  the 
birds.  When  due  allowance  is  made  for  the  kind  of 
stomach  and  absorptive  area,  apparent  differences  can 
be  reconciled.  For  example,  there  is  little  confusion 
experienced  in  comparing  acute  erosive  gastritis  or  the 
follicular  enteritis  of  an  omnivorous  intestinal  tract  (man 
or  pig),  of  a  sacculated  stomach  and  absorptive  tract  (the 
marsupial),  of  a  carnivorous  gut  (cat)  or  a  herbivorous 
compound  stomach  with  its  long  digestive  and  water- 
absorbing  surface  (cow  or  camel)  and  an  expansive 
muscular  organ  with  a  very  extensive  digestive  area 
(seal).  The  type  of  lesion  seems  the  same,  in  that  inflam- 
mation, pus,  necrosis,  granulation  tissue  and  cicatrices 
are  comparable  throughout  the  series.  The  size  of  the 
hind  gut  has  been  taken  by  Metchnikoff  as  an  indicator  of 
the  possibility  of  intoxication  by  degradation  products  of 
digestion.  He  believes  that  the  capacious  colon  of  herbiv- 
ora  and  the  short  small  one  in  carnivora  explain  the 
relatively  greater  life  in  the  latter,  because  here  less 
stagnation  and  absorption  can  take  place.  A  reference  to 
the  expected  lengths  of  life  given  before  hardly  substan- 
tiates this,  and  in  our  later  chapters  there  will  be  found 
no  strong  indication  that  animals  with  large  colons  suffer 
with  degenerative  visceral  changes  more  than  those  with 
small  ones ;  nay  even  the  reverse  may  be  found  ti-ue. 

In  regard  to  epizootics  the  behavior  of  man  and  lower 
animals  is  similar  except  perhaps  that  during  an  outbreak 
a  smaller  percentage  of  the  latter  give  evidence  of  indi- 


INTRODUCTION  39 

vidual  immunity  and  whole  groups  are  apt  to  be  carried 
off.  Occasionally  hygienic  measures  stay  the  ravages,  at 
other  times  nothing  seems  to  avail.  Fortunately  it  is 
sometimes  possible  to  sacrifice  infective  specimens  and 
remove  contagion.  We  have  had  few  serious  outbreaks, 
unless  one  might  call  our  former  heavy  infection  with 
tuberculosis  in  monkeys  an  epizootic.  The  principal  ones 
were  an  unexplained  water  fowl  disease  which  carried 
off  one  hundred  and  forty-six  birds,  an  imported  epi- 
zootic of  quail  disease  which  killed  about  the  same 
number,  a  few  cases  of  blackhead  among  wild  turkeys, 
and  a  small  group  of  cases  of  amoebic  dysentery  in 
monkeys  and  of  thrush  in  passerine  birds,  and  a  small 
number  of  tuberculous  pneumonias  in  snakes. 

Pathology  may  be  difficult  upon  an  anatomical  basis, 
but  when  we  engage  to  explain  functional  physiological 
defects  we  are  surely  embarked,  with  a  poor  compass  and 
weak  rudder,  upon  an  uncharted  sea.  One  knows,  of 
course,  that  all  animals  require  the  same  amount  of  food 
elements  per  kilo  of  body  weight,  that  man  eliminates  his 
nitrogen  as  urea  and  uric  acid,  that  monkeys  do  the  same, 
that  most  other  mammals  destroy  uric  acid  and  excrete 
allantoin,  that  birds  and  reptiles  form  uric  acid  but 
chiefly  urates,  that  there  is  an  adaptation  of  alimentary 
tract  and  diet,  that  herbivores  have  a  high  threshold  for 
carbohydrates,  that  there  is  a  variable  quantity  of 
enz\Tne  present  in  different  organs  and  in  different  ani- 
mals, that  vitamins,  whatever  they  may  be,  are  necessary 
for  the  growth  of  young  animals,  that  hormones  exist 
whereby  correlations  of  parts  are  kept  normal — ^but  these 
things,  rather  than  being  learned  thoroughly  from  ani- 
mals, have  merely  been  substantiated  by  comparisons 
with  man.  Constitutional  diseases  so-called,  from  which 
the  necessity  to  investigate  much  of  this  physiology  origi- 
nated, are  little  known  in  the  wild  animal.  Many  cases 
of  so-called  gout  have  been  encountered  and  we  have 
seen  an  instance  of  diabetes  in  a  fox,  but  more  extensive 


40     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

experience  is  needed  for  definite  practical  comparisons. 
This  applies  to  thyroid  and  pituitary  disorders  and  to  the 
vague  conditions  we  have  at  times  been  obliged  to  call 
marasmus  or  inanition. 

Some  attention  has  been  given  to  the  study  of  diets  for 
the  ^vild  specimens  of  our  Garden,  but  no  systematic 
observations  have  been  made  or  records  kept  upon  purely 
physiological  subjects.  Reference  will  be  made  at  appro- 
priate places  to  accepted  comparative  physiological  facts, 
but  our  statistics  permit  additions  to  such  knowledge  only 
in  a  limited  manner  and  in  isolated  instances.  Doctor 
Corson- White  has  very  ably  summarized  the  diet,  ali- 
mentary tract  and  physiology  of  the  zoological  groups 
with  the  pathology  as  found  in  our  records. 

A  word  might  be  added  here  as  to  the  destruction  of 
animals  by  injury  from  fighting  and  harassment  by 
others  in  the  cage.  Fighting  doubtless  causes  death, 
especially  when  males  are  together,  but  it  is  our  experi- 
ence that  in  cases  of  traumatic  death  search  should 
always  be  made  to  see  if  the  resistance  of  the  dead  animal 
had  not  been  reduced  by  some  disease.  This  is  well  illus- 
trated in  birds.  Very  frequently  a  specimen  will  come  to 
autopsy  with  its  head  feathers  plucked  out,  or  with  a  bill- 
thrust  in  the  wing  or  pelvic  region.  Such  birds  are  not 
infrequently  suffering  from  malaria,  or  heavy  intestinal 
parasitism  or  from  organic  disease  whereby  the  resist- 
ance and  self-preservatory  power  has  been  decreased. 

The  foregoing  survey  of  the  approach  to  our  subject 
reveals  the  multiplicity  of  factors  wliich  affect  the  study 
of  comparative  pathology.  No  one  of  them  can  be  entirely 
omitted,  no  one  is  without  some  effect  upon  the  origin  and 
expression  of  disease,  and  no  one  is  fully  understood. 
Yet  it  is  to  be  hoped  that  a  study  of  our  material,  accumu- 
lated under  routine  conditions  and  uninfluenced  by  any 
experimental  procedures,  will  demonstrate  the  natural 
response  of  various  zoological  groups  to  morbific 
agencies.     Perhaps   reactively  some   of  the   modifying 


INTRODUCTION  41 

conditions  may  thus  be  understood.  It  is  also  not 
unreasonable  to  expect  that  alterations  observed  as 
natural  responses  in  a  large  number  of  specimens  in 
nearly  normal  surroundings  would  serve  as  more  reliable 
guides  to  investigative  speculation  than  would  changes 
in  a  few  animals  under  artificial  technical  experimenta- 
tion. We  hope  that  the  few  facts  we  have  been  able  to 
record  may  afford  someone  a  basis  for  further  biological 
studies.  It  is  also  to  be  hoped  that  something  has  been 
learned  which  in  the  end  will  afford  an  explanation  of  the 
diseases  of  man.  Too  great  optimism  in  this  direction 
should  be  guarded  against  because  the  human  being  is 
indeed  an  animal  sui  generis  and,  from  the  standpoint  of 
normal  conditions  of  nature,  a  wild  animal. 

The  zoological  classification  found  on  pages  43-46 
was  compiled  in  1903  by  Dr.  A.  E.  Brown  on  the  basis  of 
the  British  System.  With  a  few  exceptions  the  computa- 
tions in  the  text  are  made  on  the  basis  of  zoological 
orders  since  the  number  of  specimens  in  families  is  often 
too  small  and  the  complications  of  so  many  different 
figures  would  be  confusing.  The  tables  will  be  found  to 
correspond  to  the  sequence  of  the  classification. 
Dr.  Corson- White  has,  however,  used  for  her  analysis 
the  dietary  groupings.  A  carnivore  in  her  chapter 
implies  strictly  a  meat-eater,  in  the  rest  of  the  book  one  of 
the  zoological  group  Carnivora. 

The  Laboratory  of  Comparative  Pathology  at  this 
Garden  speaks  for  the  earnest  desire  on  the  part  of  the 
Directors  to  use  the  material  to  its  fullest  extent,  and  I, 
acting  for  myself  and  my  associates,  wish  to  record  our 
appreciation  of  the  facilities  offered  to  us  for  study,  and 
for  the  broad-minded,  scientific  cooperation  the  Board  has 
always  displayed.  The  President,  Charles  B.  Penrose, 
M.D.,  Ph.D,,  LL.D.,  was  the  active  originator  of  the  plan 
whereby  this  department  was  started,  and  he  has  given  to 
it  continuously  the  support  of  his  rich  experience.  I  wish 
to  express  for  myself  the  deepest  appreciation  of  his 

4 


42     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

personal  interest  in  my  studies,  and  assistance  which  has 
been  constructive  and  stimulating.  Whether  or  not  this 
present  work  prove  useful  to  the  extent  that  is  hoped,  the 
results  from  the  Laboratory  are  such  as  to  make  the 
scientific  world  debtor  to  this  gentleman. 

It  is  a  duty,  and  a  pleasant  one,  to  record,  though 
unfortunately  in  memoriam,  my  association  with  Arthur 
Erwin  Brown,  A.M.,  Sc.D.,  Ph.D.,  C.M.L.Z.S.,  for  many 
years  the  Secretary  of  the  Society  and  Executive  Officer 
of  the  Garden.  Doctor  Brown  as  teacher  was  ever  ready 
to  help  in  the  broad  subject  of  biology,  and  I  am  proud  to 
recall  that  he  guided  me  also  as  a  friend. 

The  first  director  of  the  Laboratory  was  Courtland  Y. 
AVhite,  A.M.,  M.D.,  who  served  from  1901  to  1906,  retiring 
then  to  accept  a  position  in  the  City  Laboratory.  The 
foundation  of  the  recording  system  is  still  in  use  essen- 
tially unchanged  from  his  plan,  and  is  a  credit  to  his  fore- 
sight. Our  clerk  and  technician.  Miss  Harriet  M.  Phelps, 
has  served  the  Garden  faithfully  and  well  since  1906.  The 
condition  of  the  museum  is  very  much  due  to  her  interest 
and  watchfulness.  Thanks  and  appreciation  for  her  work 
are  felt  by  every  one,  the  author  most  of  all.  Dr.  F.  D, 
Weidman  has  been  our  first  assistant  since  1911,  and  his 
work  on  parasitology  has  been  of  the  greatest  value, 
practically  and  scientifically.  It  is  to  be  hoped  that  we 
shall  be  able  to  retain  him  indefinitely.  Dr.  E.  P.  Corson- 
White  has  in  recent  years  taken  an  assistant  position 
with  us,  armed  for  the  work  with  a  thorough  knowledge  of 
applied  organic  chemistry  and  immunology,  and  has 
already  obtained  useful  results. 


ZOOLOGICAL  CLASSIFICATION 
MAMMALIA 

PLACENTALIA 
Primates 

Simichdce  Anthropoid  apes. 

Cercopithecidce    Old  World  monkeys  (macaques,  baboons) . 

CebidcB    New    World    monkeys     (capucins,    howlers, 

spiders). 
Hapalidce   New  World  monkeys    (marmosets) . 

Lemubes 

Lemuridce     Lemurs,  Loris,  Galagos. 

Carnivoba 

Felidce    Cats. 

YiverridoB    Civets,    Genets,    Paradoxures,    Ichneumons. 

HycenidcB    Hyena. 

Canidce . .  .Dogs,  Wolves,  Foxes,  Jackalls,  Etc. 

Mustelidce    Marten,  Skunk,  Weasel,  Otter,  Badger,  Etc. 

Procyonidce    Raccoon,  Bassaris,  Coati,  Kinkajou. 

Ursidce    Bear. 

These      are 
grouped    sepa- 

Otariidce    Eared  Seal,  Sea  Lion.    J    lately   as    sub- 

PhocidcB    Common  Seal,  Walrus.    \    pET)I A     illus- 
trating   water 
carnivores. 
Insectivora 

Tenrecidce    Tenrec. 

Solenodontidce     Solenodon. 

TalpidcB    Moles,  Shrews. 

Erinaceidce    Hedgehog. 

Chiroptera 

I'teropodidcB    Fruit  Bats,  "  flying  foxes." 

VespertilionidcB Common  Bats. 

Emballonuridw   ....   Snouty  Bats,  Free-tailed  Bats. 

RODENTIA 

Sciuridce     Squirrels,  Spermophiles,  Marmots. 

Castoridce    Beaver. 

Muridce    Rats,  Mice. 

GeomyidcB    Pouched  Rats,  "  Gophers." 

DipodidcB    Jumping  Mice,  Jerboas. 

H eteromyidce Kangaroo  Rats. 

Octodontidce    Capromys,  Coypu. 

Hystricidw Porcupines. 

CMnchillidce    .*.  Viscacha,  Chinchilla. 

DasyproctidcB    Agouti,  Spotted  Cavy. 

Caviidw    Guinea-pig,    Capybara. 

Leporidce    Rabbits,  Hare. 

Proboscidea    Elephant. 

Hyracoidea     Cape  Hyrax. 


44     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Ungulata 
Perissodactyla  (odd  toed) 

Rhinocerotidce Rhinoceros.  ' 

Tapiridce    Tapir. 

Equidce Horse,  Ass. 

Artiodactyla    (even  toed) 

Bovidce  Oxen,  Antelopes,  Sheep,  Goats. 

Cervidce    Deer,  Moose,  Elk. 

Antilocapridce  Prong -horned  Antelope. 

Girafjldce    Giraffe. 

Tragulidce    Chevrotains,  Muis  Deer. 

Camelidce   Camels,  Llama. 

Hippopotamidce    Hippopotamus. 

BuidcB   Swine,  VVarthogs. 

Tayassuidcc    Peccaries. 

Sirenia    Sea-oow,  Manatee,  Durong. 

Cetacea AVliales,  Porpoises. 

Edentata 

Bradypodidce     Sloths. 

DasypodidcB     Armadillo. 

Myrmecophagidoe    ....  Ant-eaters. 

MARSUPIALIA 

Marsupialia 

Didelphyidce    Opossums. 

Dasyuridce   Dasyures,  Tasmanian  "  Devils." 

Feramelidcv    Bandicoots. 

Phascolomyidce   Wombat. 

Phalangeridce    Phalangers. 

Macropodidm Kangaroo,  Wallabies. 

MONOTREMATA 
Monotremata 

Echidnidce Echidna,  Ornithorhynchus. 

AVES  '  '-^^'^'''' 

Passeres 

Turdidcp   Thrushes,  Robins,  Etc. 

Sylviidce Warblers,  Kinglets. 

ParidcB    Titmouse. 

Troglodytidce Wrens,  Mockingbirds,  Catbird,  Etc. 

Pycnonoiidw    Bulbul. 

Crateropodidce    Babblers,  Jay -thrushes. 

Oriolidw  Griole. 

MotacilUdo}    Wagtails. 

Dicruridw    Drongos. 

Mniotiltidce   Chats,    Warblers,    "  Woodwarblers,"    Etc. 

Ccrrebidce Sugarbirds. 

Vireonidce    Vireos. 

Laniidce    Shrikes. 

Ampelidce     Waxwing. 

Hirundinidce    Martins,  Swallows. 

Meliphagidce    Honeyeaters. 

Tanagridce   Tanagers. 

Ploceidce Weavers,  Whvdah  birds,  Waxbills,  Finches, 

Etc. 


INTRODUCTION  45 

Fringillidce    Finches,     Sparrows,     Buntings,     Grosbeaks, 

Etc. 

Icteridce  Hangnests,  Troupials,  Grackles,  "  Black- 
bird," Etc. 

Sturmdw    Starlings,  Mynahs. 

Corvidce    Crows,  Jays,  Magpies,  Jackdaws. 

AlaudidcB   Larks. 

Tyrannidce Tyrans. 

Cotingidce Bellbird,  Cock-of-the-rock,  Etc. 

PiCARLE 

TJpwpcB    Hoopse. 

Trochilidce  Hummingbirds. 

Cypselidce    Swifts,  "  Chimney  Swallow." 

CaprimulgidcB   Night   hawk.   Whip-poor-will. 

CoraciidcB f-.  Roller. 

HALCYONES 

Alcedinidce     Kingfisher. 

MomotidcB    Motmots. 

BUCEROTES 

Bucerotidce    Hornbill. 

TROGONES 

Trogonidce    Trogons. 

SCANSORES 

Picidw    Woodpeckers. 

Rhamphastidw    Toucans. 

Capitonidce    Barbets. 

COCCYGES 

Citculidce    Cuckoos. 

Musophagidce    Touracous. 

PSITTACI 

Loriidce    Lories,  Lorikeets. 

Cacatuidce    Cockatoos. 

Psittacidce    Macaws,       Conures,       Amazons,       Parrots, 

Parrakeets. 
Striges 

StrigidcB Barn-owl. 

Btibonid(B All  other  owls. 

ACCIPITRES 

FalconidcB    Buzzards,  Hawks,  Falcons,  Eagles,  Etc. 

Serpentaridce Secretary  Vulture. 

Catharidce    Vultures. 

COLUMB^ 

Treronidce    Fruit  pigeons. 

Columhidte All  other  pigeons  and  doves. 

Pterocletes 

Pteroclidce    Sand  grouse. 

Gali.i 

Tetraonidw    Grouse,  Ptarmigans. 

Phasianidce    Pheasants,   Fowls,   Turkeys,   Quail,   Etc. 

Cracidce    Curassows,  Guans,  Etc. 

MegapodidcB    Brush  turkey. 


46     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Hemipodu 

TumicidcB    Hemipodes. 

FULICABI^ 

Rallidce    Rails,  Porphyries,  Gallinules,  Coots,  Etc- 

Alectobides 

Aramidce    Courlan. 

Eurypygidoe Sun  bittern. 

OruidcB  Cranes. 

CariamidcB Cariama   "  Crane." 

PsophiidcB    Trumpeters. 

LlMICOL^ 

CEdicnomidce   Thicknees. 

CharadriidcB    Plovers,    Sandpipers,    Curlews,    Woodcocks, 

Etc. 
Chionidce    Sheathbills. 

Gavle 

Lariidce    Gulls,  Terns. 

StercorariidoB    Jaeger  Gull. 

Pygopodes 

Colymhidce Loons,  Grebs. 

Alcidce    Auks,  Murrs,  Puffins. 

Impennes 

8pheniscidce Penguins. 

Steganopodes 

SulidcB    Gannets. 

PelicanidoB Pelicans. 

PhaldcrocoracidcB  .  .  .  .Cormorants. 

AnMngidce Darter    "  Water    turkeys." 

Tubinares 

Procellariidce Petrels,  Fulmars. 

Hebodiones 

Ardeidce   Herons,  Bitterns,  Egrets. 

CicomidcB Storks,  Ibises. 

Plataleiidce    Spoonbills. 

Odontogloss^ 

Phoenicopteridce    Flamingoes. 

Palamede.e 

Palamedeidce  Screamers. 

Anseres 

Anatidce   Swans,  Geese,  Ducks. 

Stbuthiones 

ApterygidcB    Kiwis,  Apteryx. 

Casuariidce    Cassowaries. 

Struthionidce Ostriches. 

Rheidce   Rheas. 

Cbtptubi 

Tinamidce       Tinamous. 


INTRODUCTION  47 

List  of  Animals  subjected  to  Autopsy  giving  the  number  of  each. 

These  Figures  are  used  to  obtain  the  percentages 

quoted  in  the  Tables  and  Text. 

MAMMALIA 

Primates 498 

Lemures 86 

Carnivora 481 

Pinnipedia 20 

Insectivora 6 

Chiroptera 5 

Rodentia 198 

Ungulata  365 

Proboscidea 3 

Hyracoidea   7 

Edentata   16 

Marsupialia 175 

Monotremata 0         1860 


AVES 

Passeres 1355 

Picarias 87 

Striges   133 

Psittaci    689 

Aceipitres    196 

Columbffi    157 

Pterocletes 0 

Galli 299 

Hemipodii  2 

Fulicariffi 35 

Alectorides 37 

Limicolas 6 

Gavipe 20 

Pygopodes 0 

Impennes 5 

Steganopodes 21 

Tubinares    0 

Herodiones 98 

Odontoglossae 6 

Palamedes   5 

Anseres    317 

Struthiones  32 

Crypturi    5         3505 

5365 


SECTION  II 
DISEASES  OF  THE  HEART 

The  heart  is  an  organ  whose  duty,  throughout  the  two 
classes  considered  in  this  study,  remains  entirely  identic, 
purely  a  physical  one  in  driving  the  blood  through  the 
corresponding  vascular  system.  The  physics  involved 
naturally  differs  between  mammals  and  birds,  but  energy 
is  derived  from  the  automatic  power  lodged  in  the  cardiac 
musculature.  Whether  this  be  neuromyogenic,  as  seems 
to  be  the  case  in  all  mammals,  or  purely  myogenic,  as  is 
probably  the  case  for  the  birds  in  which  MacKenzie  and 
Robertson  (1)  say  there  is  no  atrioventricular  bundle,  the 
result  is  the  same,  since  in  both  classes  there  is  some 
continuity  of  muscle  fibres  from  auricle  to  ventricle.  The 
gross  anatomy  varies  little  if  any  more  than  the  physi- 
ology, albeit  there  is  proportionately  greater  auricular 
capacity  in  the  mammals  than  in  the  birds,  and  indeed 
there  are  differences  within  the  classes  which  cannot  now 
be  readily  explained ;  certain  minor  variations  of  valvu- 
lar arrangement  exist,  such  as  the  absence  of  the 
membranous  right  tricuspid  in  Aves. 

When,  however,  one  considers  the  cardiac  power 
available  for  various  animals,  the  subject  becomes  one  of 
greater  breadth  and  complexity,  for  no  consistency 
obtains  even  within  families,  since  the  demand  for 
cardiac  strength  will  vary  more  with  habits  than  with 
zoological  relationships.  Thus  for  example  the  domestic 
rabbit  has  a  small  heart  volume  while  the  wild  hare  has  a 
great  one.  Although,  of  course,  the  size  of  an  organ  may 
not  be  an  absolute  measure  of  its  efficiency  (a  flea's  leg 
muscle  has  relatively  greater  power  than  a  man's),  yet 
size  is  the  only  physical  gauge  one  has  for  estimating 
nature 's  preparation  for  expected  demand.    Perhaps  this 

(1)   Br.  Med.  Jour.,  1910,  2,  1161. 
48 


DISEASES  OF  THE  HEART  49 

will  be  shown  later  when  after  discussing  the  pathological 
anatomy  of  the  heart  in  the  wild  mammals  and  birds,  we 
can  study  these  changes  in  light  of  statistics  upon  the 
relative  size  of  the  heart. 

Expression  of  Cardiac  Disease. 

The  diseases  of  this  organ  are  known  only  by  their 
physical  effects,  chiefly  by  causing  physical  or  functional 
defects  in  other  organs  and  to  a  minor  degree  by  purely 
physiological  irregularities  in  the  heart  itself  (tachy- 
cardia, arrhythmia).  All  the  latter  and  most  of  the 
former  are  subjects  discovered  by  observation  during 
life  and  unfortunately  cannot  be  included  in  the  study 
at  hand.  Both  states  are  well  known  to  the  veterinarian 
who  diagnoses  them  with  reasonable  ease  in  animals  that 
can  be  handled ;  I  saw  one  case  of  arrhythmia  in  a  monkey 
for  which  no  adequate  gross  morbid  explanation  was 
found  postmortem.  Some  of  these  functional  abnor- 
malities are  certainly  caused  by  myocardial  disease  and 
cardiac  failure  has  occurred  among  many  orders.  An 
interesting  observation  was  made  by  Plimmer  (2)  on  sev- 
eral large  birds  (ostriches,  storks,  cassowaries)  which 
apparently  died  from  this  condition ;  at  autopsy  he  found 
myocardial  degeneration,  or  epicardial  edema  or  only  a 
flabby  heart.  Lack  of  exercise  was  held  responsible  by 
this  observer.  Such  cardiac  deaths  have  probably  been 
encountered  at  this  Garden  but  we  have  accounted  them  to 
shock,  or  gastrointestinal  disease;  this  matter  will  be 
discussed  on  a  later  page. 

Coronary  Artery  Disease. 

If  a  degenerative  sclerosis  of  coronary  vessels  be  the 
cause  of  angina  pectoris  then  perhaps  paroxysms  of  this 
kind  occur,,for  we  have  seen  such  anatomical  changes  in 
the  heart  of  three  widely  separated  varieties  of  animals, 
a  Nylghaie,  a  Hamadryas  Baboon,  a  Macaque,  and  a 
Brown  Pelican;    The  history  of  these  animals  does  not 

(2)   Proc.  London  Zool.  Soc,  1907. 


50     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

register    anything    resembling    the    clinical    picture    of 
angina  pectoris  in  man,  and  they  did  not  come  to  their 
death  from  the  arterial  changes  in  the  heart  alone  since 
sufficient  other  pathology  also  existed. 
Kinds  or  Pathological  Change. 

As  an  introduction  to  the  strict  pathology  of  the  heart 
it  might  be  well  to  outline  the  headings  of  the  scheme 
upon  which  it  seems  desirable  to  study  the  subject.  It  is 
hardly  profitable  to  take  up  seriatim  the  ordinary  general 
pathological  processes  as  discussed  in  systems  of  pathol- 
ogy for  it  is  our  purpose  to  show  the  distribution  of  basic 
aberrations  from  thenormalintermsof  zoologicalposition. 
To  this  end  one  must  consider  the  response  of  the  heart 
(a)  to  damaging  influences  and  (b)  to  a  demand  for 
increased  work.  In  the  first  group  come  degenerations 
and  inflammations,  upon  which  may  succeed  an  incom- 
petency in  the  form  of  dilatation.  The  response  of  the 
normal  heart  to  any  physical  demand  greater  than 
customary  has  usually  been  thought  to  lie  in  the  direction 
of  hypertrophy,  but  on  occasion  it  has  seemed  to  be  in  the 
form  of  dilatation,  especially  if  the  strain  has  been  sud- 
den and  severe.  Starling  thinks  that  the  primary  and 
normal  reaction  of  the  heart  to  physical  strain  is  always 
dilatation.  The  idea  of  hypertrophy  must  not  be  con- 
fused with  an  understanding  of  the  relatively  large 
hearts  in  animals  whose  habits  demand  great  cardiac 
power,  for  then  it  is  their  norm  and  might  be  called 
''physiological  cardiac  hyperplasia."  I  shall  use  the 
terms  ''increased  muscle  bulk"  and  "increased  chamber 
space ' '  as  preferable  to  hypertrophy  and  dilatation ;  this 
also  focuses  attention  upon  the  two  features  of  an 
enlarged  heart. 
Degenerations  and  Inflammations. 

Degenerative  changes  in  disease  are  recorded  in  our 
system  as  amyloid,  hyaline,  fatty  metamorphosis,  gran- 
ular and  cloudy  degeneration.    While  there  is  perhaps 


DISEASES  OF  THE  HEART 

Table  1. 


51 


Table  Showing  Incidence,  in  the  Orders,  of  Degenerations  and  Inflammations, 
or  in  Other  Words  the  Response  to  Infectious  and  Toxic  Agencies. 


Heart  of 

1 

■■S 

Q 

.2 

.2 

i 

1 

CD 

i 
1 

* 

1 

21 

.t 

iJ 

Primates. . . 

8 
1 
19 
2 
5 

3 
2 
12 

4 

2 

io 

2 
2 

.',4 

2 

16 
2 

12 

14 

"9 
3 
5 

19 
10 

17 
3 

18 

li 

"6 

"8 
1 
6 

9 
3 

1 
7 
8 

13 

1 

1 

'.  '.  '. 

"1 

11 
5 

io 

;; 
2 

12 
2 

2 

6 

1 

38 
1 

54 
3 

16 

"1 

20 

7 

33 

19 
6 
1 

24 

36 

3 

28 

"3 

2 

'2 

"1 
4 

"4 

50 
11 

'5 
1 
1 

7.6 

Lemures 

1.2 

Carnivora 

11. 

Pinnipedia 

16. 

Rodentia 

8. 

Insectivora 

Chiroptera 

Proboscidea 

33. 

Hyracoidea 

Ungulata 

5.4 

Edentata 

U- 

Marsupialia 

19. 

Monotremata 

Passeres 

1.4 

Picariae 

6.7 

Striges 

.8 

Psittaci 

3.4 

Accipitres. .  .  . 

18.3 

Columbae .... 

2. 

Pterocletes.  . 

Galli 

9.3 

Hemipodii 

FiilicariEB 

8.6 

Alectorides. . 

5.4 

Limicolae. . . 

Gaviffi 

10. 

Pygopodes 

Impennes 

16. 

Steganopodes 

20. 

Herodiones 

4.1 

Palamedes 

15.7 

Struthiones .  .    .  . 

34. 

Crypturi 

Total... 

98 

118 

112 

44 

367 

8 

*  Total  is  the  number  of  individual  animals  showing  lesions,  not  the  aum  of  the  listed 
changes. 

In  this  and  subsequent  tables,  figures  in  italics  are  for  small  groups 
of  animals  coming  to  autopsy,  usually  less  than  one  hundred,  and  from 
which  percentages  may  be  misleading.  The  number  of  autopsies  upon  such 
groups  may  be  found  by  consulting  the  list  given  on  page  47. 


52     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

between  some  of  these  conditions  and  true  myocarditis  a 
matter  only  of  degree,  the  records  have  been  analyzed  as 
filed  and  perhaps  some  lessons  can  be  learned  from  the 
responses  of  the  various  zoological  orders.  In  the 
accompanying  Table  1  will  be  found  the  distribution  of 
degenerative  and  inflammatory  lesions  through  the  zoo- 
logical orders.  The  percentages  speak  for  themselves  but 
deserve  as  well  some  consideration  from  the  standpoint 
of  normal  heart  value ;  this  will  be  taken  up  later  after  the 
other  lesions  have  been  discussed. 

Endocaeditis,  Myocarditis. 

Romberg  said  in  his  classical  work  on  the  heart  that 
there  is  always  some  form  of  myocardial  disease  with 
endocarditis.  The  44  cases  of  valvular  disease  detected  at 
this  Garden  are  15  of  chronic  nature,  29  of  acute  or  sub- 
acute character.  In  the  former,  the  chronic,  9  showed 
some  grade  of  muscular  involvement,  while  21  of  the  29 
acute  cases  of  valvular  inflammation  were  accompanied 
by  myocardial  damage ;  the  percentages  are  60  for  chronic 
and  72  for  acute,  a  relation  that  would  be  expected  if  one 
credit  the  theory  that  many  valvular  inflammations  start 
at  the  root  of  the  valves,  but,  at  all  events,  they  indicate 
that  after  the  acute  stages  have  passed  the  myocardial 
damage  may  be  repaired. 

The  large  number  of  cases  of  endocarditis  among  the 
Camivora,  Accipitres,  Anseres  and  Marsupialia  is  note- 
worthy and  can  hardly  be  explained  by  other  argument 
than  a  special  vulnerability  of  this  organ  in  these  groups. 
However,  the  unusual  number  of  cases  in  our  only  native 
marsupial,  the  opossum,  seems  worthy  of  a  special  note 
since  ten  of  the  twelve  instances  in  the  order  Marsupialia 
affected  this  particular  animal.  When  seen  these  ten  cases 
were  acute  in  five  instances,  subacute  in  three  and  chronic 
in  two.  The  Streptococcus  pyogenes  was  isolated  in  three 
of  the  five  acute  cases;  bacteriology  of  the  others  was 
negative  or  not  done.     All  of  the  acute  and  one  of  the  sub- 


Fig.  1. — VEGETATIVE  AND  ULCERATIVE  ENDOCARDITIS  OF  AORTIC  VALVE.  OSTRICH 
(STRl'THIO  AISTRALIS).  ORGANISMS  ISOLATED  CORRESPONDED  CLOSELY  TO  BAC.  AFANASIEFFI- 
CHESTER. 


DISEASES  OF  THE  HEART  53 

acute  cases  were  combined  with  some  evidence  of  general 
septicemia.  The  type  of  lesion  was  in  no  way  peculiar, 
unless  the  facts  that  all  were  vegetative  when  acute  and 
markedly  deforming  when  chronic,  he  noteworthy.  In  two 
chronic  aortic  cases  the  valvular  orifice  was  almost  closed, 
yet  the  left  ventricle  could  not  be  considered  as  greatly 
hypertrophied  and  no  dilatation  existed.  In  one  acute 
aortic  and  mitral  case,  general  dilatation  existed. 
The  mitral  was  involved  nine  times,  three  times  alone, 
four  times  with  the  aortic,  once  each  with  the  tricuspid 
and  pulmonary.  Once  the  vegetations  were  hmited  to 
the  mural  endocardium.  Nine  of  these  animals  came  from 
one  enclosure  over  a  period  of  two  and  a  half  years,  dur- 
ing which  time  other  opossums  died  from  similar  bacterial 
infections  (pneumonia)  despite  repeated  cleansing  of  the 
place.  No  unusual  number  of  cases  of  this  or  similar 
kinds  occurred  elsewhere  in  the  Garden  at  this  time,  but 
it  would  seem  that  we  had  in  this  cage  a  continued  bac- 
terial infection. 

Unusual  Pericardial  Changes. 

The  appearance  of  a  stiff  gelatinous  exudate  in  the 
pericardial  sac  has  attracted  our  attention  on  ten 
occasions  (8  birds,  2  mammals).  The  substance  seems 
quite  homogeneous  and  almost  entirely  acellular.  One 
specimen  became  solid  on  heating  and  another  became 
turbid  when  put  into  Kaiserhng  's  fluid.  In  two  cases  bac- 
terial cultures  were  made ;  nothing  grew.  It  has  not  been 
associated  with  tuberculosis  or  tumors  nor  has  any  one 
pathological  lesion  more  than  another  appeared  to 
accompany  it. 

A  peculiar  lesion  occasionally  seen  in  birds  is  ''uratic 
pericarditis ' '  a  process  not  infectious  at  all,  according  to 
Plimmer,  but  due  to  renal  disease.  It  has  been  seen  here 
in  association  with  retention  of  urates  in  the  kidney,  with 
gout  of  birds,  and  apparently  quite  independent  of  any 
renal  or  constitutional  disease.    Both  layers  of  the  sac 


54     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

are  pearl  gray  or  irregularly  salted  with  a  whitish  granu- 
lar material  so  that  they  are  entirely  opaque ;  occasionally 
the  distribution  is  spotty.  The  deposit  does  not  seem  to 
penetrate  the  myocardium.  There  is  at  times  some 
involvement  of  other  serosse,  but  this  is  usually  much  less 
marked  than  around  the  heart.  It  does  not  seem  that  this 
of  itself  should  be  fatal,  but  it  has  been  the  most  decided 
pathological  factor  in  some  of  the  autopsies. 

Aside  from  pericarditic  exudates,  twenty-one  instances 
of  pericardial  effusion  have  been  encountered.  They  offer 
little  that  is  peculiar  in  etiology,  chemistry  or  cytology, 
but  as  there  has  been  some  question  of  the  position  of  the 
accumulation  of  the  fluid  in  human  beings,  it  might  be 
well  to  note  the  position  in  our  material.  It  is  recognized 
in  veterinary  medicine  that  the  cardiac  dullness  is 
increased  especially  to  the  right,  and  that  most  of  the 
fluid  will  be  on  that  side  and  posteriorly.  At  our 
autopsies  on  mammals  this  is  the  position  usuall}^  occu- 
pied by  the  fluid,  the  apex  being  covered  by  pericardium, 
unless  the  quantity  be  great  enough  to  make  the  sac  taut, 
and  this  position  is  retained  whether  the  animal  be  laid 
upon  the  one  side  or  the  other ;  nor  does  the  fluid  all  leave 
the  base  of  the  heart  when  the  body  is  placed  prone.  The 
crown  of  the  heart  is  nearly  always  well  covered.  In  birds, 
on  the  other  hand,  the  fluid  occupies  the  apical  part  of  the 
sac,  probably  due  to  the  fact  that  this  membrane  is 
attached  by  its  tip  to  the  transverse  air  sac  wall  which 
takes  the  place  of  a  diaphragm,  so  that  the  tip  of  the  heart 
is  always  free  and  the  pericardium  of  the  base  fairly 
closely  applied  to  the  epicardium.  This  holds  good  even 
for  the  birds  whose  cardiac  apex  is  normally  attached  to 
the  pericardium  by  a  fibrous  band.  ', 

Hypertrophy  and  Dilatation. 

The  response  of  the  heart  to  a  continued  demand  upon 
its  working  capacity  will,  as  already  indicated,  lead  to 
increased  muscle  bulk  or  to  larger  chamber  size.  Whether 


DISEASES  OF  THE  HEART  55 

hypertrophy  be  purely  the  building  of  a  bigger  engine  or 
be  accompanied  by,  or  due  to,  muscular  disease  as  had 
been  suggested  by  certain  authors,  was  a  question  to 
which  an  answer  was  hoped,  but  it  would  seem  that  the 
solution  is  no  nearer  than  can  be  obtained  in  human 
pathology.  Fifteen  of  the  34  cases  showed  some  degree 
of  myocardial  damage.  Some  of  the  other  cases  may  have 
been  instances  of  so-called  essential  hypertrophy, 
enlargements  due  to  hard  work  or  to  low-grade  hidden 
infection.  Aubertin  (3)  ascribes  such  cardiac  muscle 
increase  to  overwork  under  the  stimulus  of  intoxi- 
cation from  intestinal  sources  or  from  irregular 
constitutional  functions.  For  information  concerning 
this  and  pathological  enlargements  one  may  consult  the 
Table  (2)  of  Hypertrophies  and  Dilatations;  in  advance 
the  method  of  charting  must  be  known.  Since  it  is 
usually  impossible  to  decide  what  may  be  the  single 
important  factor  in  the  cardiac  disease,  all  of  the 
accredited  factors  have  been  listed  with  the  hope  that  the 
resulting  figures  would  be  significant.  Thus  an  animal 
may  have  recorded  pericarditis,  nephritis  and  arterio- 
sclerosis— ^who  shall  say  which  was  primary  or  most 
potent  in  the  cardiac  change. 

Essential  hypertrophy  is  limited  to  those  cases  for 
which  there  was  no  concomitant  pathology  that  might 
have  been  responsible  for  the  overgrowth.  There  was  one 
in  a  carnivore  (fox)  and  one  in  a  raptatory  bird  (buz- 
zard). Idiopathic  dilatations  on  the  other  hand  are  much 
more  common,  but  they  stiU  bear  a  relation  to  the 
apparent  vulnerability  of  the  heart.  Their  distribition  is 
as  follows  :  Primates  1,  Carnivora  1,  Pinnipedia  1,  Ungu- 
lata  1,  Marsupialia  4,  Anseres  1.  These  may  be  cases  such 
as  Plimmer  described,  of  cardiac  failure,  indicated  by 
dilatation,  the  result  of  inactivity.  Besides  these  special 
instances  and  the  ones  accounted  for  in  the  list,  there 
were  three  acute  dilatations  apparently  due  to  shock, 

(3)    Comptes  Rendus  Soc.  Biol,  T,  62-206. 


56     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

two  ungulates  and  one  marsupial,  probably  incidental  to 
fright  when  being  caught  by  the  keepers. 

The  association  of  secondary  dilatation  with  hyper- 
trophy is  only  evident  in  three  instances.  One  case  and 
perhaps  the  most  interesting,  is  that  in  which  the  princi- 
pal antecedent  pathology  was  thyroid  hyperplasia  and 
nephritis;  the  dilatation  was  perhaps  agonal  or  shortly 
before  the  last  struggles.  It  would  seem  that  all  of  the 
dilatations  occurred  shortly  before  death  because  long 
standing  passive  congestions  and  dropsies  of  cardiac 
origin  are  exceedingly  rare;  only  one  certain  case  is 
recorded  (carnivore). 

Let  us  now  examine  the  Table  (2)  according  to  orders 
and  then  as  to  causation.  The  Primates'  heart  is  appar- 
ently well  able  to  increase  in  size  in  response  to  increased 
work,  a  demand  most  often  made  by  pulmonary,  pleural 
and  pericardial  diseases.  Two  of  these  cases  occurred  in 
animals  suffering  with  pulmonary  tuberculosis  suffi- 
ciently extensive  to  impede  cardiac  action  while  in 
another  case  the  tuberculous  lesion  was  mild  but  a  peri- 
carditis existed.  When  the  right  hand  columns  are 
inspected  it  would  seem  that  on  occasion  dilatation  may 
occur;  one  of  the  tuberculous  pulmonary  cases  had  a 
dilated  heart.  The  slothful  lemurs  apparently  have  no 
call  upon  their  cardiac  mechanism. 

Carnivora  with  their  large  organ,  which,  it  would 
seem,  should  be  prepared  for  excess  work  either  simply 
as  a  reserve  or  as  an  inherent  ability  to  grow, 
present  in  about  equal  numbers,  hypertrophy  and  dilata- 
tion. It  is  admitted  that  there  are  within  this  order, 
genera  of  differing  habits,  but  analysis  of  the  canida?, 
felidae  and  ursida?  for  examples,  in  the  first  place,  offer  too 
few  specimens  for  conclusions  and,  secondly,  have  upon 
trial  actually  shown  nothing  definite,  so  that  we  are 
forced  to  use  the  larger  group,  the  order.  It  is  interest- 
ing to  note  that  long  continued  infection  is  in  this  order 
the  most  potent  factor  in  enlargements  of  the  heart.  Four 


DISEASES  OF  THE  HEART 

Table  2. 


57 


Table   Showing  Incidence  -per  Order  of  Hypertrophy  and  Dilatation,  and  the  Principal 
Associated  Lesions  Believed  to  Have  Etiological  Importance. 


Hypertrophy 


Primates 

Lemures 

Carnivora 

Pinnipedia 

Rodentia 

Insectivora 

Chiroptera 

Proboscidea. . .  . 
Hyracoidea .... 

Ungulata 

Edentata 

Marsupialia. .  .  . 
Monotremata .  . 
Total  Mammals 

Passeres 

Picariae 

Striges 

Psittaci 

Accipitres 

Columbse 

Pterocletes 

Galli 

Hemipodii 

Fulicariae 

Alectorides 

Limicolae 

Gaviae 

Pygopodes 

Impennes 

Steganopodes .  . 

Tubinares 

Herodiones 

Odonotoglossae . 

Palamedes 

Anseres 

Struthiones .... 

Crypturi 

Total  Birds .... 

Total 

For  meaning  of  italics  see  foot  note  Table  1 

5 


34 


2    7      7      3  13 10  6    3  15  54  0    3     10      9  14  8 


3. 

12.5 
4.5 


1.5 


58     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

of  the  ten  cases  show  myocarditis.  Nephritis  does  not 
seem  very  important  in  relation  to  cardiac  muscular 
increase,  but  occurs  with  great  frequency  in  association 
with  dilatation.  Three  of  the  cases  of  hypertrophy  were 
associated  with  thyroid  disease  and  two  of  these  showed 
dilatation  as  well.  The  general  causes  of  chamber 
distention  are  more  diverse,  and  we  see  associations 
that  do  not  appear  with  hypertrophy,  namely  arterio- 
sclerosis and  diseases  of  the  chest. 

The  rodents  seem  to  have  no  power  to  increase  muscle 
bulk,  but  a  sufficient  number  of  cases  of  dilatation  occur 
to  make  one  conclude  that  this  is  their  method  of  response 
to  unusual  strain.  Pulmonary  disease,  mostly  of  infec- 
tious nature,  and  myocardial  degenerations  are  the 
principal  causes. 

The  next  order  to  show  cardiac  enlargement  is  the 
Ungulata  where  nephritis  is  the  most  frequent  associa- 
tion with  hypertrophy  and  disease  of  the  pleura  and  peri- 
cardium with  dilatation,  or  the  reverse  of  the  factor  value 
in  the  Camivora.  These  animals,  fairly  well  prepared 
for  flight,  with  moderately  large  hearts,  seem  more  often 
to  show  dilatation  than  hypertrophy. 

Two  Edentata  (armadillo)  showed  dilatation  but 
no  hypertrophy. 

Marsupials  behave  somewhat  like  rodents  in  that  the 
heart  does  not  seem  to  increase  muscle  bulk,  but  our 
records  do  not  explain  this  clearly.  As  already  men- 
tioned four  cases  had  no  sufficient  internal  reason  for 
dilatation,  but  as  one  was  probably  the  result  of  shock 
three  only  remain  to  be  accounted  for.  Nephritis  seemed 
to  exist  in  all  three,  but  two  of  them  had  kangaroo-mycosis 
of  the  jaw  and  a  general  chronic  infection. 

If  now  our  attention  be  given  to  the  Aves  we  find  the 
highly  specialized  Passeres  and  Striges  not  represented 
and  their  closely  related  well-organized  orders  Picarife 
and  Psittaci  with  only  an  isolated  single  case.  This  is  the 
more  interesting  since  the  last  order  suffers  reasonably 


DISEASES  OF  THE  HEART  59 

often  with  arteriosclerosis.  Accipitres,  the  birds  of  pug- 
nacious habit  and  carnivorous  diet,  seem  well  able  to 
increase  their  muscle  upon  demand,  but  do  not  often 
suffer  dilatation.  Vascular  and  renal  diseases  stand  out 
most  prominently  in  the  etiology,  and  one-half  of  them 
show  myocardial  change.  The  Galli,  which  includes  both 
ground  and  flying  birds,  are  represented  but  fail  to 
exhibit  any  unusual  accompanying  disease.  Anserine 
birds  apparently  have  a  low  power  to  increase  the  size 
of  the  heart,  but  most  often  allow  it  to  dilate.  Struthi- 
ones,  large  stalking  and  rapidly  travelling  birds, 
apparently  have  a  good  margin  of  safety  in  their 
cardiac  mechanism. ) 

Summary  of  Lesions  Associated  with  Hypertrophy  and 

Dilatation. 

Analysis  of  the  associated  pathology  will  reveal  that 
among  the  mammals,  renal  disease,  chronic  infections 
and  diseases  of  the  thoracic  serosa  are  most  often  respon- 
sible for  hypertrophy,  and  that  something  over  one-third 
of  the  hearts  showed  myocardial  damage.  Among  the 
Aves  arteriosclerosis  and  renal  disease  are  most  impor- 
tant in  enlarging  the  heart;  half  of  the  cases  had 
myocarditis.  In  so  far  as  dilatation  in  mammals  is 
concerned,  renal  disease  and  acute  infections  are  decid- 
edly more  important  than  other  influences,  even  than  the 
next  in  order — chronic  infections  and  pulmonary  dis- 
eases ;  only  one-fifth  of  the  cases  had  myocardial  disease. 
Acute  infectious  disease  is  the  most  potent  cause  of  dilata- 
tion in  birds ;  only  two  of  the  eight  cases  had  degeneration 
of  the  heart  muscle. 

Comparison  of  Mammalia  and  Aves. 

If  a  comparison  of  the  incidence  of  increased  muscle 
bulk  in  the  two  classes  be  made  (4)  it  will  be  found  to 
occur  two  and  one-half  times  more  often  in  mammals, 

(4)  This  is  done  by  determining  the  percentages  of  hypertrophy  and 
dilatation  for  the  total  number  of  each  class  examined  at  autopsy. 


60     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

while  dilatation  occurs  nearly  ten  times  more  often 
among  the  mammals  than  among  the  birds.  Hypertrophy 
is  accompanied  by  myocardial  change  in  44  per  cent,  of 
the  cases,  whereas  muscular  degeneration  was  only  seen 
in  24  per  cent,  of  the  dilatations ;  this  change  is  conspicu- 
ously lacking  in  the  Primates,  Ungulates  and  Marsupials. 
The  usual  teaching  has  been  that  dilatation,  which  means 
enlargement  of  chambers  and  thinning  of  walls  or  at  least 
no  thickening  thereof,  imphed  an  inability  on  the  part  of 
the  heart  to  keep  up  with  increased  demand — a  decom- 
pensation. If  Starling  be  correct  that  dilatation  is  not  a 
degeneration  of  pump  value  but  merely  one  of  adapta- 
tions to  increased  demand,  then  this  method  is  more 
characteristic  of  mammals  than  of  birds.  There  is, 
however,  the  reserve  power  to  increase  the  muscle 
bulk  inherent  in  the  mammalian,  not  possessed  or 
needed  by  the  avian  heart.  The  large-hearted  class 
Aves  certainly  dilate  their  blood  pump  less  frequently 
than  mammals  and  indeed  have  less  cardiac  disease. 

An  analysis  of  the  incidence  of  hypertrophy  versus 
dilatation  shows  that  hypertrophying  power  resides 
in  the  Primates,  Accipitres  and  Struthiones,  their 
hearts  relatively  seldom  dilatating.  Lack  of  such  power 
and  consequent  dilatation  resides  in  Rodentia,  Ungulata, 
Marsupialia  and  Anseres.  Hypertrophying  power  lies 
therefore  chiefly  in  the  heart  of  average  size  for  its  class, 
dilatation  occurring  in  the  small  heart.     (See  page  63.) 

Avian  Hypertrophy. 

There  is  little  to  be  learned  from  the  nature  and 
anatomy  of  the  hypertrophies  and  dilatations  except  per- 
haps their  character  among  the  birds,  in  which  the 
physics  of  the  circulation  is  somewhat  peculiar.  In  this 
class  both  the  hypertrophy  and  distention  are  predomi- 
natingly left-sided,  a  state  probably  explained  by  the 
pressure  against  which  the  pump  must  work  in  flight 
because  then  the  lungs  and  the  viscera  are  somewhat 


DISEASES  OF  THE  HEART  61 

compressed  by  the  pressure  of  an  excess  of  air  in  the 
pneumatic  sacs.  At  all  events  while  concentric  hyper- 
trophy was  mentioned  once,  it  is  difficult  to  estimate  the 
degree  of  increase  in  the  right  chambers  because  they  are 
not  uncommonly  well  filled  when  diastole  occurs  at  death. 
Grober  (5)  asserts  that  the  normally  large  heart  (or  what 
I  have  called  "physiological  hyperplasia")  shows  a 
* 'hypertrophy"  of  the  right  ventricle  because  of  the  extra 
work  entailed  in  flying.  This  is  certainly  not  the  case  in 
the  material  we  have  seen  under  pathological  conditions. 
Right-sided  increase  might  be  expected  if  pulmonary  or 
serous  membrane  affections  were  prominent,  but  left- 
sided  increase,  following  arteriosclerosis  and  nephritis  is 
the  actual  finding.  The  best  examples  of  concentric 
hypertrophy  are  in  the  dogs  with  thyroid  disease  and  the 
best  examples  of  concentric  dilatation  in  ungulates 
suffering  shock.    , 

Summary. 

The  foregoing  pathological  data  can  now  be  sum- 
marized by  grouping  the  facts  under  the  headings  of 
absolute  and  relative  vulnerability  of  the  heart.  By  the 
former  is  meant  the  actual  number  and  quality  of  lesions 
in  the  various  orders,  but  here  at  once  one  comes  upon  the 
irregularity  of  examples  of  zoological  and  pathological 
character,  and  if  one  trust  entirely  to  the  percentages, 
fallacious  conclusions  might  be  reached.  Basing  judg- 
ment upon  the  incidence  of  pathological  lesions  in 
mammals  and  birds,  it  is  evident  that  the  former  has 
greater  vulnerabiUty,  as  13  is  to  6.2.  This  is  noteworthy 
as  we  shall  learn  that  the  bird  has  a  larger  and  appar- 
ently better  prepared  heart  than  the  mammal.  Attempts 
to  discover  the  order  or  kind  of  animal  having  the  great- 
est or  lowest  vulnerability  are  difficult  for  the  reason 
given  above.  Thus,  for  instance,  Pinnipedia,  Probos- 
cidea,  Edentata,  Gaviae,  Impennes,   Steganopodes,  and 

(5)   Arch.  fur.  Ges.  Physiologic,  1908,  125,  507 


62     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Strutliiones  present  the  highest  percentages  of  cardiac 
lesions,  but  the  total  specimens  examined  are  so  few  that 
these  figures  may  well  be  misleading.  (See  Tables  1 
and  2.)  If,  however,  figures  mean  anything  in  such  small 
groups,  these  are  the  animals  which  have  the  greatest 
cardiac  vulnerability.  They  have  little  in  conmaon  in 
regard  to  zoological  relationships  and  habits ;  four  of  the 
seven  orders  are  rather  slothful  and  three  are  active.  It 
is  much  better  to  limit  our  observations  to  those  orders 
from  which  sufficient  examples  have  been  subjected  to 
autopsy  and  upon  which  we  have  some  standards  for 
comparison  in  the  heart-body  weight  ratio.  It  so  happens 
that  in  the  above  seven  orders  I  was  unable  to  obtain  any 
reliable  figures  of  heart  weight.  Table  3  is  a  combination 
of  data  from  Tables  1  and  2  for  the  principal  orders 
from  which  we  have  enough  material  (at  least  one  hun- 
dred autopsies)  and  for  which  it  is  possible  to  obtain  as 
comparative  standards  figures  indicating  the  weight  of 
the  normal  heart  in  kilograms  of  body  weight; 
Table  4  gives  these  ratios  for  normal  hearts.  The 
information  about  the  weights  was  obtained  from 
some  of  our  own  figures  and  the  references  given  in 
the  footnote.  (6)  There  are  no  extensive  data 
upon  weights  and  measures  in  exact  terms,  such  as 
body  weight,  so  that  we  are  limited  to  the  numbers 
quoted  in  parentheses  besides  the  orders  in  the  table. 
The  ratios  might  be  modified  slightly  by  a  greater 
number  of  examples,  but  they  show  certain  things  by 
comparison  of  the  classes ;  in  a  rough  manner  the  heart 
ratios  correspond  to  the  pathology. 

(6)  dal  Piaz:  Papers  from  the  Department  of  Anatomy,  University  of 
California,  1912.  Bergmann:  Dissertation,  Munich,  1884.  Loer:  Arch.  f. 
die  gesamte  Physiologic,  1911,  V.  140-293.  Grober:  Arch.  f.  die  gesamte 
Physiologic,  1908,  V.  125-507.  Grober:  Deutsch  Archiv  f.  Klin.  Med.,  1907, 
V.  91,  502.  Welcher  and  Brandt:  Arch,  fiir  Anthropologie,  1903,  V.  28. 
Vierordt:  Tahellen,  1906.  Parrot:  Zoologischer  Jahresbericht,  1893. 
Hasenfeld  and  Romberg:  Arch.  f.  Exp.  Path,  und  Pharmacol,  1897,  V. 
39-333.     Joseph:  Jour.  Exp.  Med.,  1908,  V.  10-521. 


DISEASES  OF  THE  HEART 


63 


Table  3. 

Table  Containing  a  Condensation  of  the  Two  Foregoing   Tables  and  Showing 

Figures  for  Degenerations,  Hypertrophy  and  Dilatations  for 

Orders  Having  the  Largest  Number  of  Autopsies. 


Degenerations,  &c. 

Hypertrophy 

Dilatation 

7.6 
11. 

8. 

5.4 
19. 

1.4 

6.7 

.8 

3.4 

18.3 
2. 
9.3 
4.1 

15.7 

.8 
2.1 
0. 
1.2 

.6 

0. 

1.1 

0. 

0. 

4.3 

0. 

.7 
0. 

.6 

.8 

2.2 

4.2 

3. 

5. 

0. 

0. 

Striges 

0. 

Psittaci 

Accipitres 

Columbae 

GalU 

.1 
.5 
0. 

.7 

Herodiones 

Anseres 

0. 
1.5 

Table  4. 

Table  Showing  Weight  of  Normal  Heart  in  Relation  to  Body  Weight.      Number  of 
Specimens  used  to  Determine  Weight  Quoted  in  Parenthesis. 


Average  Heart  of 

Grams  per 

Kilogram  of  Body.* 

Man 

(  4) 

5.67 

Primates 

(  4) 

6.56 

Carnivora 

(  6) 

6.78 

Rodentia 

(  5) 

5. 

Ungulata 

(10) 

5.8 

Marsupialia 

(  3) 

5.1 

Average  5.82 

Passeres 

(43) 

19.8 

Picarise 

(  9) 

21.3 

Striges 

(  4) 

7.33 

Psittaci 

(  6) 

8.89 

Accipitres 

(  7) 

12.32 

Columbae 

(  4) 

14.47 

GalU 

(16) 

11.08 

Fulicarise 

(  3) 

23.82 

Limicolse 

(  2) 

8.78 

Anseres 

(14) 

11.8 

Struthiones 

(  1) 

12.7 

Average  13.84 

*  These  values  are  obtained  in  part  from  the  literature,  in  part  from  our  own  specimens. 
At  this  laboratory  the  hearts  of  animals  killed  or  dying  from  a  short  illness,  organs  showing 
no  pathological  change,  were  removed  by  cutting  at  the  base  of  the  vessels,  washed  free  of 
blood  and  weighed.    The  weight  of  the  whole  animal  was  obtained  after  death. 

For  meaning  of  italics  see  foot  note  Table  1. 

However,  there  are  many  reasons  why  great  caution 
should  be  used  in  evaluating  the  relative  size  of  the  heart. 
Welcher   showed  in   his   work   that   the   proportion   is 


64     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

greater  in  small  and  young  animals  than  in  large  and 
adult  ones.  All  the  authors  quoted  agree  that  in  birds 
and  to  less  degree  but  still  clearly  in  mammals,  there  is  a 
direct  relationship  between  the  bodily  activity  of  an 
animal  and  its  cardiac  bulk.  This  is  fairly  well  shown 
in  the  list  of  avian  heart  ratios,  but  not  so  clearly  in  the 
mammals.  What  shall  be  considered  the  most  active 
mammals — the  monkey,  perhaps,  with  his  tendency  to  be 
occupied  constantly,  yet  we  find  the  greatest  heart  bulk 
among  the  Carnivora,  animals  prepared  for  travel  and 
struggle,  and  the  smallest  among  the  Rodentia,  quiet  and 
timid  animals.  The  avian  order  showing  the  greatest 
cardiac  ratio,  the  Fulicariae,  shore  birds,  is  made  up  of 
some  quiet  hiding  varieties,  and  of  some  capable  of  very 
prolonged  flight;  the  most  constantly  active  fliers  (Pas- 
seres)  also  have  a  high  cardiac  weight  proportion.  The 
inactive  owls  have  the  smallest  heart  bulk. 

The  contrast  between  the  average  heart-to-body 
weights  of  mammals  and  birds  is  striking,  the  latter  hav- 
ing two  and  one-half  times  as  much  as  the  former, 
5.8  vs.  13.8.  Since  this  is  the  most  prominent  and  best 
supported  statement  in  the  table  of  weights,  it  may  be 
used  to  compare  mth  the  incidence  of  the  pathology  as 
seen  in  the  two  classes. 

Degenerations  and  inflammations  occur  in  mammals 
and  birds  as  9.5  is  to  5.5.(7) 

Hj^pertrophies  occur  in  mammals  and  birds  as  10.3  is 
to  4.3.(7) 

Dilatations  occur  in  mammals  and  birds  as  2.4  is 
to  ,28.(7) 

'  In  other  words,  mammals  are  much  more  susceptible 
than  birds  to  degenerative  and  inflammatory  processes, 
show  an  ability  to  increase  the  muscle  bulk  two 
and  a  half  times  as  great  and  are  liable  to  chamber  dis- 
tention nearly  ten  times  as  often.    It  might  also  be  put 

/  (7)    These   lipures   are   obtained    by   determining   mathematically   the 

percentage  of  each  feature  in  each  class  and  then  reducing  the  numbers 
to  their  lowest  value. 


DISEASES  OF  THE  HEART  65 

that  birds  cannot  or  do  not  need  to  increase  their  muscle, 
and  that  the  chamber  and  muscle  balance  is  more 
perfectly  arranged. 

While  in  the  preceding  pages  hypertrophy  has  been 
discussed  rather  from  the  standpoint  of  its  value  as  a 
compensating  and  reserve  capacity,  and  dilatation  as  a 
degenerative  or  decompensatory  process  on  the  part  of 
the  cardiac  mechanism,  it  may  be  that  dilatation  of  the 
mammalian  heart  is  the  usual  method  employed  by  the 
class  in  response  to  increased  demand.  It  seems  certain, 
however,  that  the  originally  and  normally  larger  heart, 
both  mammalian  and  avian,  more  often  uses  an  increase 
of  its  muscle  to  this  purpose. 

Hypertrophy  was  accompanied  by  myocardial  disease 
in  44  per  cent,  of  the  cases,  while  dilatation  showed  this 
change  in  only  24  per  cent.  This  supports  the  theory  that 
dilatation  is  a  normal  response  of  the  myocardium  under 
strain  and  the  belief  held  in  many  quarters  that  the  muscle 
increases  its  bulk  because  some  of  it  is  damaged. 

The  differences  between  classes  are  not  so  conspicuous 
between  orders.  However,  the  large  heart  of  the  carni- 
vores increases  both  its  muscle  and  chambers,  while  the 
small  heart  of  the  rodents  and  marsupials  more  often 
dilates.  Analysis  of  the  avian  orders  is  inconclusive  and 
somewhat  contradictory.  Let  it  suffice  to  say  that  the 
birds  which  fly  most,  with  exception  of  the  ducks,  have  a 
relatively  low  vulnerability,  and  the  soaring  carnivorous 
Accipitres  and  the  largest  birds,  Struthiones,  apparently 
have  a  high  susceptibility  to  damaging  influences  and 
enlarge  their  muscle  bulk  in  response  to  increased  work,  i 

Aneurysms  of  the  heart  are  quite  rare ;  only  two  have 
been  seen.  They  were  both  located  at  the  apex  of  the 
left  ventricle  in  birds ;  they  did  not  rupture.  Myocardial 
damage  is  evident  in  both  cases  but  the  cause  is  not  clear ; 
parasites  could  not  be  demonstrated.  Plimmer  reports  a 
case  of  cardiac  aneurysm  at  the  apex  from  infestation 
of  the  heart  muscle  by  sarcosporidia. 


SECTION  III 
DISEASES  OF  THE  BLOOD  VESSELS 

The  gross  anatomy  of  the  blood  vascular  system  is 
constructed  upon  the  same  general  scheme  throughout 
mammals  and  upon  a  comparable  basis  in  birds.  Micro- 
scopically there  is  little  variation  throughout  the  orders 
unless  it  be  in  the  relative  proportion  of  muscular  and 
connective  tissues.  The  origin  of  the  great  vessels  at 
their  cardiac  base  and  their  distribution  to  the  pul- 
monary and  to  the  greater  circulations  in  no  way  differ  in 
these  two  classes  in  that  it  always  consists  of  an  efferent 
pathway  to  the  lung  and  a  root  vessel  above  the  aortic 
orifice.  The  foiTner  has  usually  quite  a  distinct  origin 
on  the  right  side,  but  in  some  birds  the  posterior  wall  of 
the  pulmonary  artery  may  overlie  the  entire  aortic  base ; 
this,  however,  is  not  the  rule  for  birds.  The  aorta  in  most 
mammals  remains  a  separate  and  distinct  vessel  for  some 
distance,  after  which  it  gives  off  the  innominate  and  sub- 
clavians.  In  the  birds  on  the  other  hand,  the  stretch 
immediately  above  the  aortic  valve  is  usually  ballooned 
out  somewhat,  into  a  sort  of  sac  or  ampulla  from  which 
the  subclavians  and  descending  aorta  arise.  This  forms 
a  structure  of  rather  trident  shape,  the  lateral  prongs 
being  the  subclavians,  the  middle  and  posterior  being  the 
aorta  proper.  In  some  birds  the  aorta  may  have  the 
length  of  a  centimetre  or  more  then  dividing  into  the  left 
subclavian  and  right  aorta  from  which  the  right  sub- 
clavian comes  off. 

There  is  definitely  more  support  to  the  heart  and 
vascular  roots  in  mammals  than  in  birds,  in  the  latter 
class  these  structures  lying  quite  free  between  the  lateral 
air  sacs  and  well  in  front  of  the  lungs.  Nor  is  there  the 
richness  of  mediastinal  areolar  and  fatty  tissue  in  the 
winged  creatures. 


DISEASES  OF  THE  BLOOD  VESSELS  67 

The  vessels  of  mammalia  retain  a  considerable  wall 
throughout  nearly  their  entire  length.  At  first  the  wall 
is  thin  compared  to  the  calibre  of  the  vessel  while  the 
arteries  smaller  in  calibre,  have  a  heavy  wall.  In  birds 
the  arterial  stalk  at  the  heart  is  supplied  with  very  heavy 
walls,  but  after  the  second  branching  the  relation  of  wall 
to  calibre  seems  to  continue  about  the  same.  In  this  class 
the  stalk  vessels  have  wall  to  calibre  relation  of  1  to  3 
(measurements  in  2  Passeres,  1  Psittaci,  1  Accipitres) 
whereas  in  mammals  the  relation  varies  from  1  to  5  to  1 
to  7  (observations  on  2  carnivores,  1  ungulate,  2  rodents). 
In  mammals  the  consistency^  of  a  normal  artery  wall 
remains  much  the  same,  a  firm,  resilient,  yellow-white 
tissue,  quite  opaque  and  standing  open  upon  cross 
section.  In  birds  this  description  covers  the  main  stalk, 
the  aorta  in  the  abdomen  and  the  first  part  of  the  carotid 
and  iliacs.  When  these  characters  are  lost,  the  arteries 
become  semitranslucent  bluish  strands  so  that  they  are 
difficult  to  follow  in  the  muscles  of  the  neck  and  extremi- 
ties. This  is  particularly  true  in  the  Passeres,  Picariae, 
Galli  and  Columbae  while  in  the  Psittaci,  Accipitres,  and 
Anseres  the  arteries  are  distinctly  whiter  than  the  veins 
but  yet  quite  soft.  In  the  Herodiones  and  Struthiones, 
thick  walled  vessels  may  be  followed  as  far  as  the  second 
joint  in  both  extremities.  These  differences  depend  in 
part  upon  the  grosser  construction  of  the  central  arteries 
in  Aves  and  in  part  upon  the  larger  amount  of  elastic 
tissue  in  them  than  in  the  secondaries  and  smaller  vessels, 
and  than  in  comparable  mammalian  vessels. 

It  is  impracticable  to  go  into  the  minutiae  of  histology 
in  the  different  orders,  which  indeed  varies  but  little, 
although  attention  might  be  directed  to  the  facts  that  in 
all  central  vessels  the  relative  amount  of  elastica  is 
greater  than  in  smaller  ones  and  that  muscular  tissue 
seems  to  exceed  in  the  latter.  Considerable  work  has  been 
done  upon  the  amount  and  arrangement  of  muscle  bands 
in  isolated  genera,  but  no  comprehensive  data  are  at  hand 


68     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

upon  orders.  The  strands  of  muscle  do  not  seem 
arranged  so  regularly  as  in  mammals;  the  pulmonary 
artery  of  the  cat,  for  example,  has  a  muscle  arranged  like 
an  oblique  band  in  waves  or  festoons  along  the  length. 
The  mammals  as  a  class  seem  more  richly  supplied  with 
arteries  and  veins  than  do  the  birds,  and  the  square  area 
of  the  vascular  system  is  likewise  larger.  This  is  dis- 
tinctly different  from  the  amount  of  heart  bulk  as  given 
in  the  discussion  of  kilogram-heart  ratios  so  that  one 
might  say  that  the  birds  are  * '  overhearted  and 
undervesseled. ' ' 

In  so  far  as  the  physiology  of  the  two  classes  is  con- 
cerned'it  is  obvious  that  a  different  regulatory  system  is 
necessary  because,  aside  from  the  variations  of  pressure 
incidental  to  pulmonary,  muscular  and  visceral  work, 
there  remains  the  altering  pressure  within  the  air  sacs  of 
Aves,  a  force  different  under  states  of  rest,  of  running,  of 
deep  water  swimming  and  of  flying  with  or  against  the 
wind.  Part  of  the  internal  air  pressure  variation  is  cared 
for  by  the  ability  a  bird  has  to  respire  the  air  in  its  sacs 
and  bones,  but  in  prolonged  exposure  to  the  pressure 
under  water  or  during  protracted  flight  some  compen- 
satory mechanism  doubtless  exists.  This  seems  to  reside 
in  part  in  the  heavy  elastic  quality  of  the  arterial  stalk 
and  the  very  rich  venous  supply  of  the  abdomen,  includ- 
ing the  renal-portal  system  and  the  distensible  pelvic 
veins.  Just  where  the  governing  power  for  this  mech- 
anism resides  is  as  much  a  matter  of  debate  as  in  the  case 
of  the  human  being,  but  certain  researches  would  place  it 
in  the  caudate  lobe  and  pituitary  body.y 

Having  discussed  these  general  comparative  data  we 
can  now  pass  to  a  consideration  of  the  pathology  seen  at 
this  Garden.  The  subject  will  be  studied  from  the  stand- 
point of  the  vessels  as  a  system  and  the  changes  peculiar 
to  it.  Naturally  the  most  important  lesions  affect  the 
great  stalks  and  the  principal  trunks,  from  which  the 
processes  may  continue  into  the  smaller  vessels.     The 


DISEASES  OF  THE  BLOOD  VESSELS  69 

essential  alterations  are  inflammatory  and  degenerative, 
of  which  the  latter  are  by  all  odds  the  more  important. 
The  former  are  either  involvements  of  the  vessel  walls 
by  frankly  infectious  processes,  or  less  easily  proved  to 
be  bacterial  in  origin,  as  is  the  case  with  periarteritis 
nodosa.  Acute  arteritis  and  phlebitis  are  constantly 
encountered  and  present  nothing  unusual.  General  nodal 
periarteritis  has  been  seen  in  the  lower  animals,  Lupke 
having  reported  (1)  before  the  German  Pathological 
Society  a  big  outbreak  in  cows,  but  it  is  less  common  than 
among  men ;  we  have  not  discovered  it  here. 

Thromboses. 

Thrombosis  is  practically  always  a  parasitic  or  an 
infectious  process  although  at  times  considerable  difl&- 
culty  is  encountered  in  explaining  the  source  of  the 
worms  or  bacteria.j  Thus,  for  example,  the  iliac  or  femoral 
thromboses  which  are  at  the  bottom  of  intermittent 
claudication,  are  frequently  quite  vague  in  origin.  We 
have  had  one  such  case  in  a  deer  in  which  a  partly  occlud- 
ing thrombangeitis  existed  in  both  femoral  arteries  and 
veins.  Mesenteric  thrombosis,  a  serious  condition  in 
cattle  and  horses  from  infestation  with  sclerostomum  or 
strongylus,  has  not  been  proven  at  the  Garden,  but  we 
have  seen  one  case  of  numerous  thromboses  of  the  venous 
radicals  in  the  jejunal  wall  apparently  due  to  some  nema- 
tode larvae;  the  specimens  were  so  soft  by  decomposition 
that  determination  was  not  attempted.  There  occurred 
a  thrombosis  of  the  cava  and  aorta  originating  from  a 
necrotizing  cloacitis,  apparently  streptococcal  in  nature, 
in  a  Demoiselle  crane.  The  clot,  while  not  totally  occlu- 
sive, extended  nearly  as  far  as  the  heart  in  the  vein  and 
the  abdominal  aorta.  There  is  also  on  record  a  throm- 
bosis of  the  vena  cava  and  right  pulmonary  vein  in  an 
American  beaver,  harboring  Hepaticola  hepatica  in  the 
liver,  with  a  fibrosing  pneumonia  due  to  this  parasite. 

(1)    Verh.  deutsch  Path.  Gesel,  1906,  X,  149. 


70     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Another  case  in  which  parasites  seemed  to  take  a  hand 
concerned  a  common  raccoon  with  tapeworms  (sp.  ?)  in 
the  small  intestine  and  microscopically  discoverable 
parasitic  parts  in  the  lungs ;  these  organs  were  the  seat 
of  extensive  congestion  and  venous  thrombosis,  the  latter 
containing  really  enormous  numbers  of  diplococci.  The 
parasites  probably  paved  the  way  for  bacterial  invasion. 
A  frank  case  of  septic  thrombotic  aortitis  was  noted  in  a 
Rice  Grackle,  the  infectious  focus  apparently  being  a 
vegetative  *  tricuspid  "  valvuUtis.) 

Arteritis. 

In  addition  to  these  cases,  productive  inflammatory 
changes  were  discovered  five  times  affecting  vessels  in  or 
near  frank  inflammatory  processes.  The  animals  affected 
with  this  productive  process  were  three  birds,  a  rodent 
and  an  elephant.  In  the  case  of  two  birds  and  the  rodent 
the  process  was  associated  with  chronic  intestinal  lesions, 
while  in  the  elephant  it  was  found  as  an  endarteritis  oblit- 
erans in  large  vessels  of  the  lung  of  chronic  pulmonary 
tuberculosis  occurring  in  this  animal.  These  instances 
serve  as  examples  of  the  truly  productive  inflammatory 
processes  affecting  vessels  and  illustrate  the  distribution 
through  the  animal  kingdom.  Pathogenetically  there  are 
no  essential  differences,  and  histologically  they  corre- 
spond to  the  forms  seen  in  man.  Had  every  piece  of 
tissue  been  subjected  to  microscopy  wherein  such  lesions 
might  have  existed,  more  examples  might  have  been 
discovered,  but  these  processes  excite  no  peculiar 
secondary  effects  so  that  attention  is  not  dra^vn  to  them 
directly.  The  only  noteworthy  difference  between  mam- 
mals and  birds  is  the  fragile  character  of  the  clots  in  the 
latter  class.  This  is  peculiar  because  the  principal 
response  of  this  class  to  an  infectious  irritant  is  coagula- 
tion necrosis,  liquefying  enzymes  apparently  being 
absent  or  smaU  in  quantity. 


DISEASES  OF  THE  BLOOD  VESSELS 


71 


Fatty  deposits  in  the  aortic  intima  are  by  no  means 
uncommon  in  the  human  subject  and  are  encountered  at 
all  ages,  even  in  youth  at  a  time  when  progressive 
arteriosclerosis  does  not  accompany  them.  There  is  a 
belief  in  many  quarters  that  this  fat  may  be  laid  down 

Table  5. 

Tabh  Showing  the  Incidence  of  Degenerative  Arterial  Disease,  the  Percentage  in 

Animals  Subjected  to  Autopsy  and  the  Principal  Associated  Pathology. 


"o 

1 

F 

■3 

o 

CD 

is 

6^ 

^S 

Ulu 

s 

■«  S 

m 

H 

II 

2 

r 

§1 

> 

0.T3 

1^ 

^ 

•< 

S 

> 

rt 

o^ 

3 
16 
13 

.6 
3.3 
3.5 

5 

7 

3 

1 
1 

1 

? 

1 

3 

1 

5 

2 

3 

1.8 

Total 

35 

3 

2 

1.8 

.22 
2.2 

7 

7 
2 

5 

1 

2 

4 

8 

Picariae 

Psittaci 

13 
3 

1.8 
2.2 

1 

1 

6 

?, 

3 

2 

Striges...    .           .    . 

Accipitres. 

13 

6.6 

1 

4 

2 

5 

? 

4 

GaUi* 

5 

1.6 

I 

4 

2 

1 

2 

Steganopodes* 

6 

25. 

1 

3 

Herodiones 

1 

1. 

Palamedes 

1 

20. 

1 

1 

Anseres 

11 

3.4 

2 

4 

1 

4 

1 

Struthiones 

7 

22. 

1 

2 

3 

Alectorides 

2 

5.A 

1 

1 

Total 

66 

1.8 

1 

6 

15 

4 

25 

9 

n 

Grand  Total 

101 

1.8 

8 

13 

20 

5 

27 

13 

19 

*1  with  Coronary  sclerosis  only. 
**2  with  Coronary  sclerosis  only. 
For  meaning  of  italics  see  foot  note  Table  1. 

and  then  removed.  Such  deposits  are  exceedingly  rare 
in  wild  animals ;  when  they  occur  it  is  in  small  indefinite 
patches  and  not  the  bands  or  rows  as  found  in  man. 

Degenerative  Arteritis  or  Arteriosclerosis. 

Whether  or  not  it  be  exact  to  speak  of  the  more  pro- 
tracted forms  of  vascular  disease  usually  called  arterio- 
sclerosis or  atheroma  as  degenerative,  such  changes 
fonn  the  most  pronounced  features  of  the  lesions,  and 


72     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

we  have  made  such  a  separation  at  this  laboratory.  Here 
is  not  the  place  to  engage  in  the  academic  discussion  of 
the  nature  of  the  process,  but  I  A\ish  to  state  that  col- 
lectively the  changes  as  seen  in  such  lesions  in  the  lower 
animals  are  more  degenerative  than  productive,  and  that 
we  have  never  seen  true  ulcerative  atheroma  as  it  not 
uncommonly  appears  at  the  autopsy  table  in  any  large 
hospital.  This  disease  of  the  vascular  walls  has  long 
been  attributed  to  alcohol,  gout,  syphilis  and  other  such 
prolonged  intoxications  to  which  we  might  apply  the  light 
term  of  ''toxins  of  civilizations."  Too  little  credit,  or  dis- 
credit has  been  given  to  chronic  intestinal  disorders, 
overeating,  and  overdrinking  of  ordinary  fluids,  to 
entirely  incorrect  diets,  and  to  chronic  bacterial  diseases. 
Even  though  the  exact  counterpart  of  the  disease  in  man 
does  not  occur  in  lower  animals,  we  shall  see  the  probable 
association  with  food  and  with  habits,  in  a  manner 
discordant  with  former  teaching  of  the  causation  of 
the  disease. 

The  group  to  which  the  name  degenerative  arteritis 
has  been  applied  is,  as  has  already  been  indicated,  more 
productive  than  the  analogues  seen  in  the  human  being, 
but  indeed  it  is  questionable  whether  the  lesions  even  in 
the  lower  animals  are  not  more  degenerative  than  pro- 
ductive." Since,  however,  chronic  arteritis  is  always  asso- 
ciated with  damage  to  the  elastic  and  muscular  fibres  of 
the  media  as  well  as  with  fatty  change  and  overgrowth 
of  the  intima,  all  the  deforming  and  degenerative  cases 
will  be  classed  together. 

The  general  picture  in  mammals  is  one  of  diffuse 
rather  than  of  plaque-like  thickening,  but  well  outlined 
raised  or  depressed  areas  are  encountered.  In  the  aorta 
and  larger  branches  one  may  find  irregular  streaking  and 
loss  of  elasticity  with  fairly  clear,  pale  yellow  or  gray, 
flat  sections  of  distinct  opacity.  Rarely  these  may  con- 
tain calcareous  matter,  a  change  most  often  seen  in  the 
carnivores.    The  lesions  are  very  largely  limited  to  the 


KiG.   2.— ARTERIOSCLEROSIS    AND    ATHEROMA.     THORACIC   AORTA.      JACKAL   (CANTS 
AUREUS).     THIS  WAS  CONTINUED  TO  THE   MESENTERIC   AND   ILIAC   VESSELS. 


DISEASES  OF  THE  BLOOD  VESSELS  73 

aorta ;  26  or  76  per  cent,  of  the  35  cases  had  this  distri- 
bution alone.  The  arch  seemed  never  to  be  affected  alone, 
and  indeed  it  is  rather  commoner  to  find  opaque  patches 
stretching  along  the  thoracic  or  even  abdominal  portion ; 
this  is  especially  true  of  the  Ungulata. 

There  have  been  also  in  mammals  five  cases  of  mesial 
change  which  have  given  rise  to  the  picture  described  by 
Monckeberg  and  usually  entitled  by  his  name.  However, 
the  noteworthy  differences  between  the  wild  animal  and 
the  human  cases  are  the  absence  of  advanced  calcification 
in  the  media  under  the  concavities  and  the  prominence  of 
the  changes  in  the  aorta  near  the  heart  to  be  found  in 
the  former.  These  few  cases  do  not  permit  an  associa- 
tion of  the  arterial  disease  with  any  particular  pathology 
in  other  parts. 

Considered  minutely,  the  outstanding  lesion  in  the 
class  Mammalia  is  the  separation  of  the  elastic  fibres  by 
fluid  and  debris,  apparently  derived  from  the  degener- 
ated muscle  fibres,  associated  with  a  decrease  of  round 
and  elliptical  nuclei.  Globules  and  hyaline  pink-staining 
material  are  often  collected  between  split-up  elastic 
strands,  which  fibres  in  some  cases  seem  quite  numerous, 
in  others  reduced.  In  the  intima  heaping-up  of  cells  and 
fibres  is  very  moderate  in  degree  while  usually  one  finds 
only  subendothelial  edema.  When  the  process  has 
advanced  far,  the  microscopy  is  like  that  of  well 
developed  human  lesions.  Arterial  degeneration  due  to 
parasites  gives  a  different  picture  in  that  medial  degen- 
eration is  far  advanced  and  some  fibrinocellular  activity 
is  seen  upon  the  intima  when  this  tissue  remains.  When, 
however,  the  infestation  has  proceeded  to  weaken  the  wall 
sufficient  for  it  to  give  way  into  an  aneurysm,  little  or  no 
vestige  of  the  true  arterial  wall  is  left. 

In  the  Aves  the  distribution  and  anatomy  of  this 
process  present  some  differences.  The  aorta  is  as  usual 
most  conspicuously  the  seat  of  change,  but  it  is  note- 
worthy that  the  dilatation  or  ampulla  immediately  above 

6 


74     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

the  aortic  valves  and  from  which  the  main  vessels  spring, 
is  practically  always  free  of  lesions  which  are  on  the  other 
hand  most  marked  in  the  thoracic  and  abdominal  sections. 
One's  attention  is  usually  attracted  to  the  aortic  surface 
by  its  roughness  although  visibly  there  may  be  no  plaques, 
but  upon  close  inspection  a  mottled  opacity  may  be 
detected.  This  all  seems  due  in  the  few  cases  subjected  to 
tissue  section,  to  hyperplasia  of  endothelia,  with  or  with- 
out fibre  increase.  The  media  may  show  muscular  granu- 
larity or  no  change  at  all.  At  the  stage  when  plaques  are 
formed,  fairly  well  outlined,  firm  but  rather  brittle, 
raised  areas  are  detected,  seated  upon  a  distinctly  opaque 
gray  wall.  The  remainder  of  the  vessel  may  be  smooth 
and  elastic  but  sometimes,  in  the  Accipitres  for  instance, 
a  general  resistance  to  pressure  and  tension  is  found. 
Microscopically  such  a  vessel  will  show  a  media  the  seat 
of  ruptured  muscle  fibres,  split-up  or  broken  elastica  and 
some  debris,  while  the  intima  is  covered  with  active  and 
distinct  fibrocellular  exudate. 

I  have  for  comparison  divided  the  cases  into  those  in 
which  the  superficial  productive  character  was  prominent 
and  those  seemingly  entirely  a  degeneration  of  the  media, 
that  is  with  inactive  intima.  In  mammals  77  per  cent,  of 
the  cases  were  of  the  degenerative  type  while  in  birds 
50  per  cent,  were  of  this  kind^  The  exact  importance  of 
this  difference  is  not  easy  to  evaluate,  but  with 
the  facts  that  the  bird  has  a  greater  elastic  supply 
for  its  large  vessels  and  a  greater  wall-to-calibre  ratio, 
it  is  interesting.  The  aorta  alone  was  affected  in  half  of 
the  birds,  the  remainder  showing  lesions  in  the  carotids 
and  f  emorals.  , 

The  tendeiicy  for  the  media  to  degenerate  would  lay 
the  basis  for  concavities  on  the  intimal  surfaces  after  the 
type  seen  in  Monckeberg  's  sclerosis.  A  number  of  cases 
of  this  variety  have  been  encountered,  but  instead  of 
being  better  developed  in  the  vessels  of  extremities  as  in 
man  they  have  presented  more  definite  pictures  in  the 


Fig.   3.— arteriosclerosis.   ATHEROMA  AND   ANEURYSMAL   DILATATIONS  IN 
THORACIC   AORTA.      WILD   CAT   (KELIS   RUFKUS). 


DISEASES  OF  THE  BLOOD  VESSELS  75 

aorta  and  pulmonary  vessels.  Examples  will  be  quoted 
under  the  discussion  of  the  various  orders. 

Primates  are  not  often  affected  with  degenerative 
arterial  disease,  two  of  the  instances  observed  showing 
this  change  confined  to  the  coronary  vessels.  As  might  be 
expected  the  myocardium  in  both  was  affected,  and  in  one 
animal  had  a  definite  concentric  hypertrophy.  A  very 
interesting  case  was  encountered  in  a  Lion-tailed 
Macaque  {Macacus  silensis  ).  His  heart  showed  distinct 
fibrosis  of  the  conducting  pathways  from  auricle  to 
ventricle  and  of  the  papillary  muscles.  No  atheroma 
was  present  in  the  aorta,  but  in  the  pulmonary  distinct 
sacculations  of  the  Monckeberg  type  were  found.  Mesial 
degeneration  was  apparently  responsible,  but  no  calcifica- 
tion had  occurred.    Death  was  due  to  acute  gastritis. 

Carnivora  present  about  half  the  cases  seen  in 
mammals  and  16  per  cent,  of  the  total.  Five  of  the  six- 
teen cases  owe  their  origin  to  parasitic  arteritis  and  were 
combined  with  aneurysms.  As  will  be  noted  by  consulta- 
tion of  the  list  there  is  no  outstanding  accompanying 
pathology,  a  fact  which  makes  parasites  more  important. 
One  case  of  mesial  degeneration,  resembling  the 
Monckeberg  type  was  observed  in  the  thoracic  aorta  of 
a  bear. 

Ungulata  are  generously  represented,  thirteen  cases 
being  recorded  distributed  rather  unevenly  between  the 
odd-toed  (1  or  8.5  per  cent.)  and  even-toed  groups  (12  or 
2.9  per  cent.).  The  single  case  in  the  first  group  occurred 
in  a  Zebra  {Equus  burchelli)  wherein  was  found  about 
the  middle  of  the  thoracic  aorta  a  diffuse  thickening  of 
media  and  intima  in  a  circular  plaque  approximately  two 
centimetres  in  diameter;  it  was  by  no  means  so  well 
developed  as  similar  lesions  in  the  even-toed  ungulates. 
This  recalls  the  expression  of  doubt  by  ZinserUng  as  to 
the  occurrence  in  the  horse  of  arteriosclerosis  similar  to 
that  in  human  beings. 


76     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

The  lesions  in  the  Artiodactyla  are  both  in  plaques 
and  diffuse,  the  aorta  and  its  branches  sometimes  being 
quite  wrinkled  but  beset  with  firm  elevations  with  and 
without  calcification.  These  changes  are  fairly  definite 
and,  although  they  never  attain  the  development  seen  in 
man,  resemble  the  stage  of  wrinkling  and  roughness  in 
the  preulcerative  stage  of  the  human  analogue.  Two  of 
the  cases  were  associated  with  dilatations  of  the  mesen- 
teric vessels  and  with  periarteritis,  a  picture  strongly 
suggesting  parasitic  infestation;  in  one  case  ineffectual 
search  was  made,  in  the  other  no  record  is  made  of  the 
parasites.  Renal  disease  occurred  only  twice  and  myo- 
cardial damage  only  once.  Chronic  pulmonary  disease, 
present  thrice,  took  the  form  of  tuberculosis  twice  and 
pleural  adhesions  with  atelectasis  once. 

Marsupialia  present  three  quite  interesting  cases,  a 
Tasmanian  Devil  (Sarcophilus  ur sinus)  and  two  Kanga- 
roos (Macropus).  The  first  showed  distinct  sacculations 
in  the  ascending  aorta,  suggestive  of  Monckeberg's 
sclerosis  but  equally  resembling  several  small  or  incom- 
plete aneurysms.  The  underlying  vessel  was  opaque  and 
stiff,  continuing  so  to  the  middle  of  the  thoracic  portion. 
The  other  marsupials  showed  distinct  mesial  damage 
with  early  calcification  and  a  roughened  intima ;  once  the 
pulmonary  artery  was  involved. 

It  will  be  noticed  that  the  orders  Rodentia  and 
Lemures  are  missing  from  those  showing  arterial  disease 
although  we  have  a  reasonable  number  of  autopsies 
upon  them. 

Passeres  are  hardly  good  exponents  of  vascular 
disease,  an  interesting  thing  in  view  of  their  large  heart, 
heavy  vessels,  and  flying  habits.  It  is,  however,  striking 
that  two  of  the  three  birds  of  this  group  had  aneurysms  of 
the  aorta,  one  of  which  ruptured  just  above  the  origin, 
partly  into  and  partly  outside  the  pericardium.  Despite 
several  microscopical  sections  we  were  unable  to  find  the 
tiny  ends  of  the  breach  and  any  evidence  of  parasites.    In 


Fig.  4.— arteriosclerosis  IN  AORTA.  OTTER  (I.LTRA  CANADENSIS).  THERE  IS 
RELATIVELY  LITTLE  INTIMAL  CHANGE,  THE  MEDIA  BEING  FIBROTIC  AND  ALLOWING 
THE  INTIMA  TO    BE   DEPRESSED    IN   SMALL  CONCAVITIES. 


DISEASES  OF  THE  BLOOD  VESSELS  77 

the  other  case  a  vegetative  growth  occurred  on  the  intima 
near  the  dilatation. 

The  PicariaB  were  represented  by  a  Hornbill  and  a 
Toucan.  The  former  presented  roughened  yellowish  ele- 
vations for  two  centimetres  above  the  aortic  valve ;  this 
seemed  the  only  involvement.  The  Toucan  had  a 
few  small  scattered  but  deep  yellow  plaques  in  the 
same  location. 

Psittaci,  although  failing  to  be  accredited  with  a  high 
percentage  of  arterial  disease,  nevertheless  present  some 
striking  and  interesting  changes.  In  the  first  place,  the 
central  vessels  are  not  so  conspicuously  the  principal  seat 
of  atheroma  as  is  the  case  in  many  other  orders,  and  the 
lesions  are  not  so  productive.  It  is  usual  to  find  flat  areas 
of  opacity,  perceptible  as  easily  by  section  through  the 
wall  as  by  holding  up  the  opened  vessel  to  the  light, 
the  seat  of  the  density  being  in  the  media.  This  can  be 
followed  into  the  wing  arteries  and,  upon  microscopic 
section,  these  smaller  vessels  will  show  mesial  degener- 
ation, thus  being  comparable  to  arteriocapillary  fibrosis 
of  man.  However  frequent  this  picture  may  be,  there  are 
also  instances  of  overgrowth  in  the  intima,  prominences 
over  opacities  or  raised  plaques  at  the  points  of  branching, 
lesions  which  correspond  to  the  activity  of  the  inner  coat. 
It  is  interesting  that  renal  disease,  chronic  pulmonary 
and  general  infections  occur  in  a  goodly  proportion  of 
these  birds. 

Striges  is  an  order  of  little  importance.  The  lesions 
in  the  two  cases  consisted  of  rather  prominent  plaques  in 
aorta  and  large  branches. 

Accipitres  stand  out  as  giving  the  greatest  percentage 
of  any  order  of  which  we  have  had  a  fair  number  upon 
which  to  make  comparisons.  Their  arterial  lesions  are 
frequently  accompanied  by  renal,  myocardial  and  valvu- 
lar disease.  Mesial  and  intimal  alterations  are  about 
equal  in  degree,  irregular  patches  going  on  to  softening 
without  ulceration,  and  early  calcification  being  quite 
prominent.  The  sickle  at  the  branching  of  the  renals  is  a 


78     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

favorite  site  of  deposit.  Like  the  parrots  their  arterial 
damage  is  not  confined  to  the  aorta,  but  may  be  found  in 
the  carotids,  femorals  or  small  ^\dng  arteries.  The  most 
common  situation  is,  however,  in  the  lower  thoracic  and 
renal  regions. 

Gain,  represented  by  five  specimens,  seem  to  have 
their  vascular  disease  accompanied  very  often  by  myo- 
cardial and  infectious  disease.  Their  arterial  pathology 
consists  of  raised  gray  patches  in  the  aorta  only. 

Steganopodes  give  the  highest  percentage  among 
avian  orders,  but  this  must  be  held  sub  judice  because  of 
the  small  number  of  specimens  subjected  to  autopsy. 
Renal  disease  occurred  in  three  cases.  Their  lesions  are 
in  the  form  of  yellow  opaque  streakings  in  the  aorta  and 
its  branches,  in  one  case  following  the  carotid  half  way  up 
the  neck.  Plaques  are  not  common,  and  when  they  occur 
are  streaky  and  illy  outlined. 

Herodiones,  represented  only  by  a  heron,  are  neg- 
ligible. This  bird  showed  a  diffuse  thickening  with  early 
thrombotic  deposits  attached  to  the  intima. 

■  Palamedes  are  represented  by  a  Screamer  which 
showed  around  the  orifices  of  the  renal  arteries  an  early 
fatty  deposit  and  mesial  opacity. 

Anseres  present  a  considerable  number  of  cases  which 
are  accompanied  by  cardiac,  renal  and  general  pathology. 
The  character  of  the  lesions  is  like  that  of  the  Accipitres 
and  the  distribution  differs  in  only  one  particular.  In 
four  of  these  eleven  cases  the  elevations  or  opaque  areas 
were  limited  entirely  to  the  stretch  of  aorta  which  might 
be  compared  to  the  arch  in  mammalia,  that  is  the  part 
with  which  the  right  subclavian  is  in  closest  contact  and 
which  bends  almost  directly  backward  to  become  the 
thoracic  aorta.  This  excludes  the  ampulla  just  above 
the  aortic  valves.  There  seems  no  real  reason  for  this 
and  it  may  be  accidental. 

Struthiones,  with  seven  specimens  having  arterial 
disease  in  the  great  vessels   alone,   seem   to   have  no 


4    '   ■> 


KIG.    5.— AORTIC   ARTERIOSCLEROSIS.      SARUS   CRANE   (GRUS  ANTIGONE).      THE  LESIONS 
ARE  ELEVATED  AND   IRREGULAR. 


DISEASES  OF  THE  BLOOD  VESSELS  79 

especial  characters  unless  these  be  in  the  heavy  furrow- 
ing and  stiffness  of  the  wall,  with  opaque,  elevated, 
indefinite  patches,  seen  mostly  in  the  descending  aorta; 
once  a  long  tough  and  partly  brittle  stretch  was  found  in 
the  carotid. 

Alectorides,  with  a  relatively  high  percentage,  present 
irregularly  outlined  fatty  and  finely  granular  patches 
in  the  intima  of  the  lower  aorta  and  abdominal  vessels, 
and,  in  two  cases,  as  far  as  the  vessels  of  the  lower 
extremities  could  be  followed. 

The  orders  Columbae  and  Fulicariae  are  missing  from 
the  list  of  Aves  having  arterial  disease,  yet  a  reasonable 
number  of  autopsy  records  are  at  hand. 

Summary  on  Arterioscleeosis. 

Having  discussed  the  orders  separately,  a  review  of 
the  whole  situation  is  desirable.  Chronic  arteritis,  or  as 
it  is  usually  called  arteriosclerosis,  is  common  to  very 
many  zoological  orders,  and  its  principal  lesions  are 
comparable  throughout  the  two  classes  under  considera- 
tion. A  statement  as  to  its  incidence  would  best  be  made 
by  adopting  an  arbitrary  number  of  observations  as  the 
desired  minimum  upon  which  to  draw  conclusions,  and  I 
shall  adopt  one  hundred  as  such  a  figure.  Accepting  this 
as  reasonable,  a  review  of  the  table  indicates  that  the  order 
of  percentage  incidence  is :  Accipitres  6.6,  Ungulata  3.5, 
Anseres  3.4,  Carnivora  3.3,  Striges  2.2,  Psittaci  1.8,  Mar- 
supialia  1.8,  Galli  1.6,  Primates  0.6,  Passeres  0.22;  the 
other  orders  have  less  than  one  hundred  specimens  each. 
There  is  no  doubt  that  carnivorous  birds  have  the  highest 
incidence  of  chronic  arterial  disease.  Next  in  order  come 
three  varieties  with  nearly  equal  incidence,  the  ungulates, 
anserine  birds  and  carnivorous  mammals.  These  orders 
have  little  in  common  unless  it  be  that  in  nature  they  are 
often  engaged  in  prolonged  or  strenuous  effort,  as  in 
fight  or  flight.  We  possess  no  measurement  of  their 
vascular  supply  but  by  consultation  of  the  table  giving 


80     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

heart  weights  (page  63)  it  will  be  found  that  three  of 
them  have  values  below  that  of  the  class  in  which  they 
belong ;  the  Carnivora  alone  have  a  greater  heart-to-body 
ratio  than  the  average  for  its  class  Mammalia.  Nor  do 
these  orders  have  any  direct  dietetic  relationship.  The 
expected  longevity  of  these  groups  does  not  permit  one 
to  discover  any  reason  for  arterial  changes  except  per- 
haps that  they  have  a  reasonably  good  viability  under 
park  conditions,  and  therefore  many  have  a  longer  oppor- 
tunity to  develop  vascular  disease.  It  so  happens, 
however,  that  the  first  four  groups  are  the  most  likely  to 
suffer  from  gastrointestinal  inflammation,  of  dietetic  or 
bacterial  origin. 

It  is  interesting,  but  not  easily  explicable  that  the 
orders  of  great  activity.  Primates  and  Passeres,  are  at 
the  end  of  the  list ;  their  food  is  very  largely  carbohydrate 
in  character.  Just  why  Lemures,  Rodentia  and  ColumbaB 
should  be  missing  is  not  quite  clear,  because  orders  of 
comparative  habits  and  food  are  included. 

A  review  of  the  concomitant  pathology  reveals  the 
fact  that  nephritis  more  often  accompanies  these  proc- 
esses than  any  other  single  condition.  Among  the  chronic 
infectious  disease  in  the  table  is  included  chronic  enter- 
itis ;  this  group  falls  well  behind  the  renal  diseases.  The 
relatively  small  number  of  cases  of  valvulitis  speaks 
rather  against  an  active  infectious  origin  of  the  vascu- 
lar lesions. 

Aneurysms. 

Aneurysms  have  been  observed  all  over  the  world  and 
in  nearly  all  the  larger  orders.  The  London  Garden  has 
had  an  unusually  large  number  to  report,  the  most 
striking  being  that  described  by  Seligman  in  the  1906 
Report  of  the  Society,  in  a  tiger  thirteen  years  in  cap- 
tivity which  had  fourteen  sacculations  from  pea  to  plum 
size  scattered  along  the  aorta.  Even  with  the  number  of 
cases  on  record  and  those  collected  here  it  would  be 


Fig,  6. — PARASITIC  ANEURYSM  IN  THORACIC  AORTA  PARADOXURE  (PARADOXURUS  LEU- 
COMYSTAX).  PARTLY  SACCULAR,  PARTLY  DISSECTING  ANEURYSM  WITH  OPEN  THROMBOSIS 
AS  INDICATED  BY  THE  CLASS  ROD.  PIECES  OF  WORM  FOUND  IN  WALL.  COULD  NOT  OBTAIN 
WHOLE  SPECIMEN,      SPECIES   UNDETERMINED. 


DISEASES  OF  THE  BLOOD  VESSELS  81 

unwise  to  draw  deductions  as  to  their  incidence  or  as  to 
the  possibilities  of  vascular  dilatation  in  any  given  order. 
Horses  have  aneurysms  occasionally,  cows  and  dogs 
rarely,  according  to  Rievel.  I  can,  however,  state  that 
there  has  not  occurred  in  our  experience  a  large  growing 
pulsating  aneurysm  in  the  aortic  arch  region  comparable 
to  the  condition  so  well  known  in  man.  The  literature  to 
which  we  have  had  access  gives  a  definite  impression  that 
parasitism  of  vessel  walls  is  the  most  important  factor 
in  the  causation  of  ectasia,  and  that  simple  non-parasitic 
arteriosclerosis  is  relatively  unimportant.  Two  of  our 
seven  cases  seem  to  have  been  free  of  parasites  but  the 
notes  cannot  entirely  assure  one  of  this.  The  distribution 
of  cases  at  this  Garden  is  found  in  Table  5. 

A  dilatation  of  the  first  part  of  the  arch  in  a  seal  to  a 
size  which  might  be  described  as  an  aneurysm  caused  us 
to  make  such  a  diagnosis,  correctly  enough  from  the  size 
and  shape  of  the  vessel  but  possibly  worthy  of  reconsid- 
eration in  light  of  the  fact  that  no  damage  to  the  wall  was 
found.  At  the  heart  and  in  the  descending  arch  the 
diameter  measured  4-5  cm.,  while  the  first  part  of  the 
aorta  measured  7.5  cm.  This  great  irregularity  in  width 
could  not  be  found  in  other  seals  albeit  this  section  of  the 
arch  is  usually  a  trifle  larger  than  its  origin  and  descend- 
ing portion.  The  cava  in  seals  is  also  large,  but  in  this 
particular  animal  it  measured  6  cm.  across  at  the  fiver 
where  there  is  a  normal  dilatation.  These  two  spaces  are 
looked  upon  as  normal  reservoirs  for  blood  during 
diving,  but  the  case  in  question  seemed  to  have  excessive 
''aneurysmoid"  enlargements  without  mural  disease. 
London  reports  an  aneurysm  of  the  aorta  in  a  seal.  (2) 

Aneurysms  are  not  so  common  in  birds,  the  incidence 
being  in  comparison  with  mammals  as  1  to  2.2  Two  seats 
are  prominent  for  their  development :  the  sinuses  above 
the  aortic  valves  and  the  first  part  of  the  subclavian  ves- 
sels ;  two  of  the  six  cases  occupied  the  first  position,  two 

(2)   Proc.  London  Zool.  Soc,  1916. 


82     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

the  second,  while  one  other  lay  in  the  arch  of  the  aorta, 
the  last  in  its  descending  thoracic  portion.  Those 
developing-  over  the  valves  seem  to  arise  from  simple 
degenerative  arteritis;  those  that  occupy  the  other  loca- 
tions are  apt  to  be  surrounded  by  plaques  on  the  intima. 
Aneurysms  in  birds  reveal  by  microscopy  some  trace  of 
all  the  vascular  coats  and  seem  not  to  construct  an  adven- 
titia  from  surrounding  areolar  tissue.  The  veins  have 
presented  no  peculiar  pathology,  except  in  tumors  which 
will  be  taken  up  later.  A  Derby 's  Tyran  showed  a  phle- 
bitis and  periphlebitis  of  the  left  subclavian  vein,  of 
mycotic  nature,  which  led  to  death  by  rupture 
and  hemorrhage. 


SECTION  IV 
DISEASES  OF  THE  BLOOD  AND  BONE  MARROW 

The  production  and  physiology  of  the  circulating*  blood 
seem  closely  similar  in  the  two  classes  under  considera- 
tion, although  the  anatomy  is  not  the  same  in  birds  and 
mammals,  variations  also  occurring  within  the  latter 
group.  Pathological  responses  follow  comparable  lines  in 
that  hemolyzing  agencies,  be  they  hemosporidia,  absorp- 
tions from  metazoan  parasites  or  bacterial  toxins,  produce 
a  reaction  in  erythropoietic  centres,  and  positively  chemo- 
tactic  viruses  call  forth  increases  in  the  colorless 
elements.  We  have  also  observed  a  decrease  of  leuco- 
cytes in  an  Orang  Utan  suffering  from  influenza,  a  finding 
analogous  to  that  in  the  human  attack.  There  is,  how- 
ever, a  much  less  ready  response  on  the  part  of  birds  to 
any  leucocyte-stimulating  influence,  in  this  class  the 
mononuclears  seeming  to  bear  much  of  the  burden 
assumed  by  the  myeloid  cells  of  Mammalia  or  at  least 
appearing  on  the  stage  very  quickly  so  that  any  increase 
of  the  latter  is  overshadowed  by  them.  Perhaps  this 
apparent  difference  may  be  further  explained  by  the 
greater  number  of  colorless  blood  cells,  structures  which 
might  be  called  the  principal  secondary  defences  of  the 
body  and  constantly  at  the  disposal  of  the  organism, 
normally  present  in  the  birds'  blood;  they  amount  to 
25,000  per  cubic  millimetre  in  birds,  while  in  the  mammals 
very  few  varieties  have  half  this  number.  On  the  accom- 
panying Table  (6)  will  be  found  a  few  differential  leuco- 
cyte counts  now  known  to  us. 

It  would  seem,  from  a  general  observation  of  simple 
and  infected  wounds  and  from  a  few  blood  counts,  that 
the  response  of  leucocytes  in  the  lower  animals  is  greater 
than  in  monkeys  and  man.    The  ease  with  which  animals 

83 


84     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 


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DISEASES  OF  BLOOD  AND  BONE  MARROW    85 


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86     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 


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DISEASES  OF  BLOOD  AND  BONE  MARROW    87 

endure  a  wound  and  the  rapid  local  pus  formation  about 
an  infection  speak  for  an  easy  mobilization  of  their 
cellular  defenders ;  their  connective  tissue  elements  seem 
equally  well  brought  into  play.  In  so  far  as  birds  are  con- 
cerned perhaps  the  normally  large  number  of  leucocytes 
and  the  participation  of  local  tissue  cells  in  response  to 
irritation  is  a  preparatory  protective  mechanism  because 
of  their  relatively  small  amount  of  bone  marrow  which 
may  not  be  able  to  mobilize  new  cells  rapidly ;  many  of  the 
birds,  notably  those  prepared  for  long  flight,  have  much  of 
their  osseous  system  given  over  to  air  space.  The  number 
of  red  blood  cells  is  also  greater  in  Mammalia,  wliich 
show  a  variation  from  4,000,000  per  cubic  millimetre  in 
some  small  genera  to  12,000,000  per  cubic  millimetre  in 
some  ungulates,  while  birds  vary  from  2-5,000,000  per 
cubic  millimetre.  Despite  these  fundamental  differences 
in  the  classes,  pathological  changes  of  anemia,  leucocy- 
tosis  and  leucemia  are  comparable;  polycythemia  in 
lower  animals  is  unknown  to  me  but  may  of  course  occur. ) 

Anemia. 

As  in  human  pathology  this  condition  may  be 
divided  into  the  group  that  follows  some  disease  which 
damages  the  red  blood  cells  or  their  source,  called  sec- 
ondary, and  those  cases  not  preceded  by  such  a  condition, 
called  primary.  )  [Formerly  this  latter  group,  known  as 
progressive  pernicious  anemia,  was  copiously  repre- 
sented, but  study  has  discovered  that  worms,  inorganic 
poisons  and  infections  can  produce  a  picture  of  grave 
anemia  so  that  the  formerly  large  group  has  dwindled. 
We  now  conceive  a  primary  anemia  to  be  one  without 
discoverable  responsible  antecedent  pathology,  therefore 
a  disease  of  the  bone  marrow  itself.  There  is  one  variety, 
hemolytic  anemia,  which  seems  to  be  an  intoxication  of 
the  bone  marrow  with  solution  of  red  cells,  but  the 
affected  tissue  puts  up  some  struggle  against  the  poison. 
In  another  primarj^  the  so-called  aplastic  anemia,  no 


88     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

activity  at  all  is  shown  by  the  marrow,  no  young  cells 
appearing  in  the  circulation.  Clorosis,  or  green  sickness 
of  young  persons,  is  a  primary  anemia  and  presents  itself 
as  a  moderate  cell  reduction  with  a  disproportionately 
low  hemoglobin  percentage.  As  a  disease  entity  this  does 
not  occur  in  the  lower  animals,  so  far  as  I  am  aware,  but  a 
very  few  hemoglobin  estimations  and  a  reference  to  the 
literature  would  indicate  that  well  marked  hemoglobin- 
anemia  does  occur. 

In  so  far  as  the  pathology  of  anemia  is  concerned  we  are 
obliged  usually  to  judge  by  the  appearance  of  the  blood 
and  tissues,  the  yellowish  pallor  of  the  mucous  membranes, 
the  condition  of  the  marrow  and  the  amount  of  pigment ; 
severe  rapid  cases  show  hemorrhages  and  prolonged 
cases  have  fatty  degeneration  of  the  parenchymatous 
organs.  For  a  decision  of  the  primary  or  secondary 
nature  we  must  judge  the  accompanying  pathology  and 
the  condition  of  the  bone  marrow. 

Secondaey  Anemia. 

Secondary  anemia  can  be  laid  in  general  to  insanitary 
housing  or  inappropriate  diet  over  a  long  period,  to 
chronic  bacterial  infection  of  low  grade,  to  the  action  of 
blood  parasites  or  those  of  the  bowel  which  either  suck 
blood  or  elaborate  an  absorbable  toxin,  or  to  single  great 
or  repeated  small  hemorrhages.  We  shall  now  consider 
the  cases  at  the  Garden.  Perhaps  many  other  animals 
have  had  a  substandard  blood,  but  these  are  the  cases  in 
which  the  gross  appearance  attracted  close  scrutiny  in 
this  direction.  London  has  had  much  anemia,  probably 
from  their  reported  heavy  infestation  with  parasites,  but 
this  factor  has  with  us  apparently  played  a  small  role  in 
the  production  of  anemia. 

In  Mammalia. 

In  so  far  as  the  Primates  are  concerned  the  one  out- 
standing cause  of  anemia  is  degenerative  disease  of  the 
osseous  system.    In  both  rickets  and  osteomalacia  there 


DISEASES  OF  BLOOD  AND  BONE  MARROW    89 

is  an  irregular  hyperplasia  of  the  marrow,  which  is 
usually  more  marked  in  the  latter.  In  osteomalacia  one 
finds  irregular  areas  of  congestion  or  even  hemorrhage 
besides  masses  of  a  gelatinous  fatty  tissue  while  scattered 
about  are  pink  spots  where  the  marrow  is  better  pre- 
served. In  rickets,  on  the  other  hand,  the  tissue  is  more 
uniformly  congested  and  less  sharply  separated  from  the 
endosteal  osseoid  material  or  the  irregular  epiphyseal 
spongiosa.  The  fibroid  or  osteoid  growth  of  osteomalacia 
seems  to  be  fairly  well  differentiated  from  the  marrow 
tissue  although  it  may  send  strands  into  the  canal  and 
across  the  spongy  area.  Histologically  there  is  not  a  dis- 
tinct difference  in  the  appearances  nor  do  they  differ 
from  the  human  analogue.  In  those  cases  which  develop 
late  in  life  the  red  cell  centres  are  very  few  in  number  but 
usually  active.  In  the  blood,  one  finds  a  few  nucleated 
and  stippled  cells,  but  not  much  change  in  size  and  shape 
of  the  erythrocytes. 

It  does  not  seem  that  the  anemia  can  be  the  cause  of 
death,  for  among  our  thirty-nine  cases  of  osteomalacia 
and  rickets,  the  prosectors  have  thought  it  of  sufficient 
importance  to  record  in  the  diagnoses  but  eleven  times. 
There  are  usually  complications  of  pneumonitis  or  enter- 
itis to  finish  the  animal  before  the  poverty  of  the  blood 
will  do  so,  and  our  records  show  only  a  Black  Spider 
Monkey  {Ateles  ater),  a  Silky  Marmoset  (Leontocehus 
rosalia)  and  a  macaque  (sp.f)  with  osteomalacia  and 
grave  secondary  anemia.  It  would  seem,  however,  that 
hemoglobin  anemia  must  exist,  for,  despite  one  record  of 
40  per  cent.,  Fleischl,  no  excess  of  pigment  deposit  is 
noted  in  the  spleen,  liver  or  marrow. 

Carnivora  have  shown  a  moderate  number  of  diseases 
of  the  skeleton  but  the  occurrence  of  a  marrow  involve- 
ment seems  less  frank,  although  the  anatomical  changes 
are  similar.  However,  there  are  three  grave  secondary 
anemias  recorded  in  eleven  carnivores  suffering  from 
osteomalacia  and  rickets.     Another  prime  cause  of  low 


90     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

blood  value  in  this  order  is  gastrointestinal  inflammation ; 
in  seventeen  cases  of  anemia,  of  secondary  nature,  five 
showed  gastroenteritis  of  severe  grade  or  protracted 
character.  Perhaps  the  most  prolific  single  cause  of  this 
blood  change  is  parasitism,  six  of  the  seventeen  cases 
showing  infestation,  five  of  which  are  nematodes  and  two 
cestodes,  one  sho^ving  both.  The  details  of  these  are 
worth  recording.  Two  young  Jungle  Kittens  [Fells 
cliaus)  from  the  same  litter  died  of  enteritis  mth  a  notice- 
able anemia :  they  harbored  in  their  upper  small  intestine 
ascarids,  and  one  of  them  had  a  few  hookworms  (sp.?). 
A  Kinkajou  (Potos  caudivolvulvus)  died  from  a  general 
mild  infection,  emanating  from  a  pneumonia  perhaps, 
and  showed  a  heavy  infestation  with  taenia  (sp.?).  An 
American  Wild  Cat  (Felis  ruffus)  died  from  acute  catar- 
rhal enteritis  and  anemia;  the  parasitological  findings 
included  Filaria  f  asciata,  adults  in  abdominal  and  gluteal 
muscles,  larvae  in  the  blood;  Dibothriocephalis  felis, 
Ascaris  mystax  and  Uncinaria  canina  in  the  small  intes- 
tines. 'V\Tiile  the  blood  was  thin  and  pale  and  some 
pigmentation  existed,  the  condition  could  not  be  called  a 
picture  of  grave  anemia.  A  noteworthy  finding  was  the 
deep  pigmentation  of  almost  the  entire  intestinal  wall. 
Ascaris  mystax  was  found  in  an  under-sized  inbred  gray 
wolf,  killed  because  of  poor  coat;  there  was  a  marked 
anemia  and  atrophy  of  the  skin.  An  Ocelot  {Felis  cJiihigo- 
nazon)  gave  a  picture  of  anemia  due  to  uncinaiiasis,  but  is 
not  so  instructive  as  the  following.  A  Swift  Fox  {Canis 
velox)  exhibited  clearly  a  case  of  progressive  secondary 
anemia  from  uncinaria,  and  the  history  is  worthy  of  a 
brief  recital. 

Muscles  atrophic,  greenish  black  over  abdomen.  Fat  absent.  Lung 
is  blotched  by  darker  red  markings  where  parenchyma  contains  distinct 
excess  of  frothy  pink  watery  fluid.  No  fluid  or  adhesions  in  pericardium. 
Heart  is  contracted,  and  muscle  is  pale  yellow  and  firm.  The  only 
abnormality  consists  of  slight  yellowing  of  musculature.  The  abdomen 
shows  great  omentum  firmly  adherent  to  fundus  of  bladder.  No  fluid 
or  other  adhesions  in  abdomen.     The  liver  is  normal  in  size,  smooth  in 


DISEASES  OF  BLOOD  AND  BONE  MARROW    91 

surface  and  has  sharp  edges,  is  friable  and  bright  orange  yellow.  The 
section  surface  is  glistening,  smooth  and  diy.  Organ  is  poor  in  blood 
content,  greasy  and  breaks  easily.  Bladder  is  large,  contents  fluid  gT'een 
bile;  duct  patulous.  Spleen  normal.  Kidney  is  small  and  smooth, 
diminished  in  bulk,  normal  location,  smooth  surface,  and  pale  yellowish 
gray,  consistency,  soft,  flabby.  Ureters  normal.  Mouth  and  teeth 
normal.  Stomach  contains  scanty  brown  fluid.  Duodenum — Mucosa  bile- 
stained,  contained  a  solitary  nematode  worm.  In  its  lower  portion  it 
becomes  filled  with  a  blackish  red  fluid,  and  its  mucosa  becomes  studded 
by  heavily  outlined  punctate  hemorrhages  whose  positions  are  best  seen 
through  serosa.  Jejunum  similar  in  condition  to  duodenum  and  contains 
four  small  nematode  worms.  At  one  point,  i.e.,  where  the  worm  is  located 
at  beginning  of  ileum  the  blood  staining  of  mucosa  is  strictly  in  neighbor- 
hood of  the  worm.  Capillaries  nowhere  congested.  Ileum  contains  slight 
amount  of  brownish  black  material,  mucosa  normal.  Large  intestine  and 
rectum  normal.  Pancreas  normal.  Lymphatics  normal.  This  is  a  case  of 
death  by  anemia  as  result  of  bites  of  hookworms.  There  were  certainly 
more  than  four  worms  present  antemortem  since  no  males  were  found, 
and  this  may  be  explained  by  a  possible  diarrhoea  which  has  flushed  them 
out.  This  idea  is  borne  out  by  empty  condition  of  gastrointestinal  tract. 
Animal  Parasites — 'Uncinaria  canina.  The  four  small  nematode  worms 
above  mentioned  conform  in  all  respects  to  the  given  anatomical  points 
of  uncinaria.  All  four  specimens  are  perfectly  formed  females.  The  loca- 
tion of  the  hooks  was  easily  determined  and  established  as  being  in  the 
most  dorsal  portion  of  buccal  cavity,  and  as  projecting  forward  and 
ventrally  in  two  groups  of  three  each  in  same  manner  as  described  here- 
tofore in  similar  infestations  in  foxes.  The  large  nematode  worm  found 
in  duodenum  is,  from  its  possession  of  three  lips  and  its  general  form,  an 
ascaris.  The  mustache,  however,  which  is  so  commonly  seen  in  ascaris 
worms  from  this  region  is  absent.  Postmortem  blood  of  heart  shows  red 
cells  granulated  and  almost  worthless  for  histological  study.  Nucleated 
reds,  however,  are  absent  but  other  points  valuable  in  settling  the  ques- 
tion of  anemia  cannot  be  determined. 

Lung. — There  are  no  abnormalities  in  supporting  tissue.  Alveolar 
walls  are  markedly  congested.  Bronchi  normal  and  show  no  trace  of 
larval  hookworm  infestation.  Air  sacs  contain  many  red  blood  cells 
together  with  a  few  heart-failure-cells.     Congestion  of  lung. 

Liver. — No  excess  of  fibrous  tissue.  Finer  details  of  structure  can- 
not be  made  out  owing  to  advanced  autolytic  changes.  The  only  possible 
pathological  changes  consist  of  localized  areas  where  liver  cells  have 
fused  to  form  notable  masses  of  pink  granular  material  suggesting 
local  necroses.     Autolysis. 

Intestines. — Four  sections  are  present  and  all  show  essentially  the 
same  character  of  changes.  Interstitial  tissue  between  muscular  tissue 
and  submueosa  loosely  arranged  as  though  separated  by  edema. 

On  luminal  side  of  muscular  mucosa  is  a  distinct  zone  of  striking 
tawny  yellow  color  Avith  hematoxylon-eosin  combination.     This  zone  is 


92      DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

smooth  homogeneous  and  contains  a  few  spindle  and  stellate  cells 
with  no  capillaries  and  with  little  or  no  fibrillation.  It  abuts 
upon  the  fundi  of  the  crypts.  Interstitial  tissue  of  mucosa  is  loose 
and  infiltrated  with  round  and  spindle  cells  in  its  deeper  portions. 
Here  it  is  also  congested  but  congestion  is  most  marked  toward  lumen 
where  masses  of  free  blood  cells  occur  in  interstitial  tissue,  Avithin 
lumina  of  crypts  and  within  lumen  of  intestine  itself;  no  parasites  or 
ova  are  found.  Epithelium  of  crypts  has  granular  cytoplasm;  goblet 
cell  formation  frequently  seen;  cilia  well  preserved.  Edema;  Subacute 
catarrhal  enteritis  Hemorrhage. 

Six  instances  of  nephritis,  four  parenchymatous  and 
two  diffuse,  are  recorded  in  the  total  of  seventeen  cases 
of  secondary  anemia  in  carnivores.  Except  in  the  skeletal 
disease  the  marrow  is  very  mildly  affected,  some  edema 
and  reddening  grossly  and  moderate  hyperplasia 
minutely,  being  the  only  noteworthy  changes. 

The  next  order  is  that  of  the  Ungulata  wherein  we 
have  found  but  two  cases  of  frank  anemia,  an  Isabelline 
Gazelle  {Gazella  Isabella)  and  an  Aoudad  {Ovis  tragela- 
plius).  The  former  had  several  lesions  of  different 
etiology  and  nature  so  that  an  impoverishment  of  the 
blood  is  not  astonishing:  parasitic  (?)  cyst  in  lung, 
chronic  infective  arthritis,  calcareous  tuberculosis, 
congestion  and  edema  of  lungs,  osteomalacia,  and 
osteofibroma  of  maxilla.  The  condition  of  the  auodad 
was  too  indefinite  to  permit  conclusions. 

The  marsupials  are  represented  by  two  common 
Opossums  {Didelphys  virginiana)  and  a  Rufous  Rat 
Kangaroo  {Aepyprymnus  rufescens),  two  of  which  suf- 
fered also  from  rickets.  The  condition  of  the  bone 
marrow  was  unfortunately  not  recorded,  but  in  other 
rachitic  marsupials  this  tissue  follows  the  changes  seen  in 
other  orders.  One  opossum  had  a  hypertrophic  gastritis 
with  numerous  Physaloptera  turgida,  a  worm  frequently 
associated  with  chronic  thickening  of  the  mucosa ;  there 
was  also  an  early  portal  hepatic  cirrhosis  with  enlarge- 
ment of  the  spleen. 


DISEASES  OF  BLOOD  AND  BONE  MARROW    93 

Among  the  Rodentia  we  have  had  anemias  in  a 
Beechy's  Gopher  {Citellus  grammurus  beecheyi)  and  a 
Southern  {Sciurus  niger  niger)  and  Western  Fox  Squir- 
rel (Sciurus  rufiventer).  The  first  had  a  myeloma  also, 
and  will  be  discussed  later.  One  of  the  squirrels  had 
osteomalacia,  while  the  other  suffered  with  diarrhoea  and 
showed  hydropic  degeneration  of  the  kidneys,  conditions 
probably  due  to  acute  intoxication. 

In  Aves. 

The  class  Aves  is  represented  by  the  orders  Passeres, 
Psittaci,  Columbae,  Herodiones,  Gaviae,  Picarige,  Striges, 
Galli  and  Accipitres ;  the  first  four  are  well  represented, 
but  in  the  other  orders  only  one  or  two  cases  have  occur- 
red. The  causes  of  anemia  in  birds  are  essentially  those 
discussed  for  mammals  with  the  provision  that  greater 
attention  must  be  paid  to  parasites,  particularly  those  of 
the  blood.  Several  slightly  varying  protozoa  inhabit  the 
blood  corpuscles  of  birds,  and  numerous  embiyos  may 
circulate  after  they  escape  from  a  parent  lying  in  some 
organ  or  tissue.  The  role  of  blood  parasites,  intra-  or 
extracorpuscular,  in  the  cause  of  death  or  of  anemia  is, 
however,  somewhat  paradoxical.  Plimmer  seems  to 
credit  a  heavy  infestation  with  great  value  in  the  cause 
of  death.  In  the  human  being  an  infestation  of  one  cell 
in  a  hundred  is  a  fair  grade  of  malaria ;  such  a  relation  is 
apparently  common  in  birds,  and  we  have  repeatedly 
seen  a  much  heavier  seeding  while  Plimmer  reports  as 
many  as  70  per  cent,  of  the  erythrocytes  to  be  carriers  of 
hemogregarines  (he  has  seen  92  per  cent,  in  reptiles).) 
Can  then  the  effect  upon  hemic  function  and  vital  resist- 
ance be  great  ?  It  has  been  our  practice  to  interpret  the 
finding  of  circulating  protozoa  or  of  larval  metazoa  as 
merely  reducing  the  resistance  of  the  birds  so  that  they 
succumb  more  readily  to  incorrect  food,  strange  environ- 
ment or  infection. 


94     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

It  is  perhaps  well  to  show  the  state  of  our  records  in 
the  Passeres  by  a  table. 

Passeres — Secondary  anemia  associated   with 

intestinal  or  visceral  parasites   11 

blood   parasites    9 

intestinal  inflammation    6 

skeletal  diseases  and  chronic  infection   (osseous)  3 

nephritis    4 

miscellaneous  and  unassociated  anemias   8 

(entries  in  this  line  not  included  under  any 
other  heading) 

Total  cases 37 

In  the  first  group,  two  of  the  birds  showed  cestodes, 
one  a  taenia,  the  other  not  examined  for  identification; 
three  showed  coiled  filaria  in  the  air  sacs,  two,  tropido- 
cerca  in  the  proventricular  wall  and  five  had  coiled 
filaria  in  the  serosa  of  the  stomach.  The  second  group 
was  infested  five  times  with  Halteridium  and  five  times 
with  embryo  filariae.  Anemia  was  associated  with  gastro- 
intestinal inflammation  alone  only  once,  the  remaining 
five  cases  having  other  finding  of  greater  significance.  The 
three  birds  of  the  next  group  concerned  one  with  tubercu- 
losis, one  with  mycosis  and  one  with  a  long  continued 
abscess./'  It  seemed  worthwhile  to  separate  four  cases  of 
anemia  in  which  nephritis  was  a  prominent  association, 
in  three  indeed  being  the  only  other  diagnosis.  While 
it  is  impossible  to  state  that  either  is  dependent  upon  the 
other,  and  they  may  of  course  be  coincidental,  it  is  never- 
theless noteworthy  that  such  an  obscure  relation  occurs 
here  as  well  as  in  human  pathology.  In  one  of  these  birds 
seen  recently  there  was  in  all  probability  a  distinct  hemo- 
globin anemia  suggested  by  jaundice,  pallor  of  the  tissues, 
absence  of  pigmentations  and  the  finding  of  large  pale 
erythrocytes  in  the  heart  blood.  The  last  group  is  a 
mixed  one  including  some  birds  in  which  only  anemia  was 
diagnosed,  others  with  prolonged  hemorrhages,  two 
tumors,  intestinal  sand,  congestion  of  the  lungs  and 
the  like. 

The  parrots  and  their  relatives  are  represented  by 
nine  specimens,  among  which  two  had  proventricular 


DISEASES  OF  BLOOD  AND  BONE  MARROW     95 

spiroptera,  two  had  long  standing  tuberculosis  and  two 
had  osteomalacia.  The  notes  of  the  other  three  are  not 
sufl&cient  to  warrant  deductions. 

Herodiones  showed  eight  cases  of  anemia,  five  herons, 
one  bittern  and  two  storks.  Parasites  are  noted  in  only 
three  examples,  herons,  and  it  is  noteworthy  that  these 
all  had  flukes  in  the  proventricle  or  intestine;  one  also 
had  ascarids  in  the  proventricle.  Two  of  this  order  suf- 
fered with  long  standing  inflammation  following  bone 
injuries.  Perhaps  the  outstanding  features  of  this  order 
are  the  erythrocytic  picture  and  the  condition  of  the 
spleen.  The  red  blood  cells  seem  very  fragile  or  soft,  for 
one  often  encounters  in  their  fresh  or  stained  preparation 
vacuoles  or  rifts  in  the  protoplasm  surrounding  the 
nucleus.  At  first  we  thought  these  were  hemosporidia, 
but  repeated  attempts  at  their  coloration  and  the  absence 
of  pigment  granules  seem  to  warrant  an  assumption  that 
they  are  artefacts.  In  five  of  the  seven  instances  there  is 
very  definite  evidence  of  present  or  past  activity  of  the 
spleen.  We  have  not  always  considered  it  sufficiently 
prominent  to  call  it  a  splenitis,  but  follicular  activity  is 
commonly  discoverable,  and  two  cases  of  definite  fibrosis 
are  recorded.  The  sun  bittern  {Eurypyga  Jielias)  showed 
a  chronic  interstitial  nephritis  in  the  atrophic  stage.  No 
other  of  the  wading  birds  showed  secondary  anemia. 
There  are  seven  cases  among  the  pigeons  (Columbae) 
where  anemic  tissues  attracted  our  attention.  Three  were 
associated  with  osteomalacia,  in  one  of  which  the  marrow 
picture  was  that  of  an  aplastic  form  being  everywhere 
pale  and  flabby  without  cells  under  the  microscope ;  it  is 
further  interesting  in  this  case  that  there  was  a  distinct 
but  ineffectual  attempt  at  bony  regeneration  by  the  peri- 
osteum. In  another  case,  this  time  brought  to  death  by  an 
enteritis  and  cloudy  swelling  of  the  viscera,  the  marrow 
was  hyperplastic  and  red,  there  being  activity  in  the  basic 
staining  areas  of  the  head  and  in  the  shafts.  (Notes  of 
the  third  case  scanty.)    None  of  the  seven  cases  seems  to 


96     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

have  been  associated  with  animal  parasitism;  one  had 
tuberculosis.  The  other  cases  are  obscure  and  not 
definitely  connected  mth  other  pathology. 

Ten  more  cases  of  anemia  were  scattered  among  seven 
orders.  There  is  nothing  striking  or  even  individual 
about  them  worthy  of  special  mention. 

Summary  of  Secondary  Anemia. 

A  review  of  our  records  shows  that  among  5365 
animal  autopsies  we  have  recorded  anemia  of  probable 
secondarj^  character  in  122  instances,  53  (2.8  per  cent.) 
mammals  and  69  (1.9  per  cent.)  birds.  The  orders  repre- 
sented, with  the  percentage  for  the  order,  are  Primates, 

25  or  5  per  cent. ;  Carnivora,  18  or  3.7  per  cent. ;  Ungulata, 
3  or  .8  per  cent.;  Marsupialia,  4  or  2.2  per  cent.; 
Rodentia,  3  or  1.5  per  cent. ;  Passeres,  37  or  2.7  per  cent. ; 
Psittaci,  9  or  1.3  per  cent. ;  Herodiones,  8  or  8.  per  cent. ; 
Columba?,  7  or  4.7  per  cent. ;  Picariae,  2  or  2.3  per  cent. ; 
Striges,  Galii,  Gaviae  and  Accipitres,  each  one  case.  A 
consideration  of  their  associated  pathology  reveals  the 
fact  that  four  changes  are  prominently  associated  with 
secondary  anemia,  to  wit :  gastrointestinal  inflammation, 

26  times  (15  mammals  and  11  birds) ;  parasitism,  29  times 
(7  mammals  and  22  birds) ;  osteomalacia,  24  times  (18 
mammals  and  6  birds),  and  nephritis,  18  times  (12  mam- 
mals and  6  birds) ;  a  few  of  these  cases  overlap,  but  this  is 
rather  the  exception  than  the  rule,  and  this  does  not 
militate  against  the  importance  of  the  connection  with 
anemia.  It  will  be  noted  that  practically  all  the  impor- 
tant orders  of  animals  are  represented,  including  species 
from  all  over  the  globe.  There  is,  however,  no  especial 
relation  of  anemia  to  the  kind  of  diet  or  digestive  tract. 
Conclusions  as  to  the  meaning  of  these  figures  of 
incidence  are  hardly  justifiable.  In  so  far  as  the  blood 
picture  is  concerned  we  can  only  record  the  qualitative 
appearance  and  the  effect  upon  tissues.  It  cannot  be 
stated  that  to  external  observation  a  secondary  anemia 


DISEASES  OF  BLOOD  AND  BONE  MARROW    97 

presents  any  distingiushing  features  that  a  specimen  in 
poor  condition  may  not  exhibit.  The  monkeys  formerly 
dying  of  tuberculosis  had  not  infrequently  pale  buccal 
mucosa  and  skin  around  the  eyes,  but  upon  exam- 
ination of  their  viscera,  blood  or  marrow  the  quality  of 
their  blood  could  not  be  called  greatly  substandard.  Slide 
smears  of  secondary  anemia  in  many  specimens  would 
occasionally  show  stippling  or  a  moderate  number  of 
nucleated  cells  mth  anisocytosis  and  poikilocytosis. 
This  is  much  more  frankly  exhibited  in  the  Aves,  wherein 
displaced  karyolytic  or  pyknotic  nuclei  are  very  common. 
Mention  has  been  made  of  the  rifts  in  the  protoplasm, 
seen  in  Herodiones,  and  this  has  been  observed  in  other 
orders.  Perhaps  the  most  striking  change  is  the  increase 
of  young  erythrocytes  and  of  thrombocytes  in  the  winged 
creatures.  The  nucleus  of  the  former  reminds  one  of  that 
of  the  human  plasma  cell. 

The  condition  of  the  bone  marrow  corresponds  with 
fair  accuracy  to  that  which  one  is  accustomed  to  see  in 
the  human  being.  Certainly  this  holds  good  for  the 
mammals,  while  among  the  birds,  the  few  observations 
upon  which  we  feel  like  relying  indicate  a  nodular 
erythropoiesis  of  rather  striking  character.  In  the  areas 
of  reddening  as  seen  grossly  there  will  be  found  under 
the  microscope  an  orderly  arrangement  of  large  red  cells 
with  loose  chromatic  nuclei  about  a  very  much  larger  cell 
of  the  same  type,  apparently  the  primary  erythroblast. 
Outside  of  this  group,  red  cells  such  as  appear  in  the 
circulating  fluid,  are  rather  irregularly  distributed  in  a 
marginal  zone.  I  have  seen  small  areas  like  this  in 
apparently  normal  marrow,  but  the  central  grouping 
was  not  so  large  as  in  the  anemic  cases ;  it  thus  appears 
that  we  probably  have  the  anatomy  of  erythropoiesis. 

The  deposition  of  pigment  in  the  birds  is  in  much 
coarser  granules  than  among  the  mammals,  in  the  former 
case  large  masses  sometimes  obscuring  several  liver  cells 


98     DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

or  apparently  blocking  a  lymphatic  sinus;  the  Kupffer 
cells  do  not  seem  to  be  heavily  laden. 

The  extrameduUary  formation  of  blood  cells  has  been 
a  matter  of  considerable  interest  and  study  in  the  human 
being,  and  as  far  as  it  concerns  the  circulating  mono- 
nuclears, the  general  opinion  seems  to  be  that  such  a  his- 
togenesis exists.  A  decision  in  the  negative  is  perhaps 
reached  by  the  majority  in  the  case  of  erj^thropoiesis,  and 
as  far  as  my  observations  go,  this  holds  for  all  mammals. 
It  seems  worth  while,  however,  to  record  an  occasional 
finding  in  some  birds,  especially  anemic  ones,  which  may 
be  of  importance  in  their  erythropoiesis.  The  adult  red 
cell  is  a  clearly  formed  ellipse  with  a  distinct,  deeply 
stained,  sharply  outlined  nucleus  of  a  shape  correspond- 
ing to  that  of  the  whole  cell.  Young  red  cells  have  a  more 
nearly  circular  outline  but  almost  truly  circular  nucleus, 
the  short  diameter  being  at  least  proportionately  greater 
than  is  the  corresponding  short  diameter  of  the  whole 
cell;  this  is  also  the  nucleus  whose  internal  structure 
resembles  that  of  the  human  plasma  cell.  Groups  of  such 
cells  have  been  seen  in  the  interstices  of  the  liver,  some- 
times as  many  as  twelve,  in  a  rather  orderly  formation. 
An  excess  seems  at  times  visible  in  the  spleen  but  not  in 
orderly  arrangement.  Observations  are  under  way 
toward  determining  the  relation  of  this  finding  to  the 
amount  of  marrow,  the  condition  of  the  blood  and  the 
habits  of  the  bird. 

Peimaky  Anemia. 

As  already  specified  primary  anemia  is  apparently 
causeless,  aside  from  the  assumption  that  it  is  a  disease 
of  the  marrow  itself.  Since  there  are  only  four  cases, 
representing  three  orders  and  they  cannot  be  grouped  as 
could  the  secondary  variety,  the  individual  instances  will 
be  discussed  separately. 

Ring  tailed  Bassaris  {Bassariscus  astutus).  Adult  died  after  two 
weeks'  stay  in  the  Garden  with  a  history  of  general  failure  of  condi- 


DISEASES  OF  BLOOD  AND  BONE  MARROW    99 

tion.  The  diagnosis  at  autopsy  was  primary  anemia,  fatty  degeneration 
of  the  liver,  hemon-hages  in  intestines  and  spleen,  hyperplastic  bone 
marrow.  The  external  appearance  is  of  general  good  condition,  fair 
skin,  mucous  membranes  pale.  Lungs  collapsed,  and  gray  red.  There 
are  several  small  hemorrhages  scattered  irregularly  throughout  respira- 
tory tissue.  Lymph  Nodes — small,  soft  mottled  gray-red.  Pericardium 
had  slight  excess  clear  fluid,  and  no  adhesions.  Heart  normal  in  size, 
and  of  pale  brown  color.  The  liver  of  normal  size,  smooth  surface 
and  sharp  edges,  of  a  pale  brown  color,  soft  and  friable.  Has  indis- 
tinct markings  like  yellow  brown  mottlings  on  section  surface.  Gall- 
bladder contains  some  viscid  brown  bile.  Spleen,  normal  or  slightly 
less  in  size,  consistency  firm,  capsule  pale  pearl  gray,  apparently  not 
thickened.  There  are  numerous  small  hemorrhagic  spots  on  section 
surface.  Interlying  pulp  is  homogeneous  deep  red.  Follicles  not  visible. 
Kidney,  normal  in  size  and  shape.  Trabeculse  faint.  Capsule  smooth, 
strips  easily,  smooth  surface,  and  brown.  Consistency  soft.  Medulla 
prominently  striated,  cortex  homogeneous  salmon  pink.  Stomach  con- 
tains a  little  glistening  mucus.  Mucosa  pale,  flat  yellowish,  slightly 
opaque.  There  is  a  recent  clot  lying  in  some  mucus  just  above  pyloric 
valve.  There  is,  however,  no  open  vessel  nearby.  From  pylorus  to 
anus  lumen  contains  some  rather  fresh  smeared  out  or  slightly  clotted 
blood  and  mixed  in  with  mucus.  Mucosa  is  flat  translucent,  submucosa 
slightly  injected  in  a  mosaic  fashion,  otherwise  gut  wall  is  negative. 
No  recognizable  food  present.  Follicles  not  visible.  Mesentery  glands 
small,  soft,  pale  yellow.  Bones  seem  entirely  normal.  Marrow  of 
long  bones  is  firm,  bloody.  Marrow  of  ribs  also  deep  red.  Blood 
in  intestinal  tract  is  probably  a  recent  slow  oozing  from  intestinal 
Avails,  and  Avas  probably  the  last  straw.  Cause  of  this  anemia  could 
not  be  determined.  Blood  preparation  not  made  because  it  was  too 
long  after  death.  Liver  shows  moderate  fatty  infiltration  of  marginal 
areas.  Pigment  is  scarce,  only  a  few  granules  being  present  in  the 
Kupffer  cells,  not  more  than  is  often  seen  Avithout  marked  anemia. 
There  is  a  slight  increase  in  interstitial  nuclei  but  not  in  fibres.  No 
obstruction  or  increase  of  bile  ducts. 

Kidney. — Very  mild  SAvelling  of  tubular  epithelium  but  no  exuda- 
tive processes.  Glomeruli  show  a  few  vacuoles  but  capsular  space  is 
negative  to  pigment.  Bone  marroAV  (Femur)  fairly  cellular  in  con- 
struction, but  fat  Avell  mixed.  Cellular  areas  well  arranged,  active, 
most  of  cells  are  small  members  of  the  larger  mononuclear  variety. 
Small  lymphocytes  abundantly  represented.  Most  of  the  larger  cells 
are  non-granular,  Avith  centrally  placed  nucleus.  Megakaryocytes  fairly 
numerous,  nuclei  seem  closely  jammed  into  centre.  No  recognizable 
certain  nucleated  red  blood  cells,  moderately  number  stippled  cells, 
few  adult  red  cells.  Eosinophiles  and  basophiles  quite  feAV.  Pigment 
small  quantity. 

This  is  a  case  of  primary  anemia  of  moderate  severity 
and  short  duration,  and  probably  of  hemolytic  character 


100   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

if  one  may  judge  by  the  bone  marrow,  although  excessive 
pigmentation  of  the  liver  and  kidneys  was  not  found. 
Unfortunately  the  spleen  was  not  minutely  studied,  nor 
was  the  central  nervous  system  investigated.  Atrophy  of 
the  intestinal  tract  did  not  exist. 

Two  cases  occurred  among  the  monkeys,  but  one 
example  will  answer,  since  the  two  were  essentially 
the  same. 

The  case  to  be  cited  was  that  of  a  Japanese  Macaque  {Macaciis 
fuscatiis).  2  Young,  weight  three  pounds  two  ounces,  exhibited  in  the 
Garden  about  four  months,  and  apparently  in  good  shape  until  two 
weeks  before  death  when  it  rapidly  became  emaciated. 

Diagnosis. — Aplastic  anemia,  chronic  atrophic  gastritis.  Atrophy 
of  heart  muscle,  fibrosis  of  liver,  slight  local  cloudy  swelling  of  liver. 
Perilobular  diffuse  nephritis  (subcapsular  type).  Congestion  of  spleen. 
Fibrillar  fibrosis  of  spleen.  Hemosiderin  pigmentation  of  spleen.  Local 
amyloid  infiltration  of  spleen.  Calcareous  infiltration  in  medulla 
of  adrenal. 

Coat  only  fair,  body  emaciated.  Pale  muscles,  fat  scant^^  Respira- 
toiy  tract  normal  throughout  save  for  slight  emphysema.  The  Pericar- 
dium showed  no  fluid  or  adhesions.  Epicardium  glistening  and  slightly 
thickened.  Heart  pale  in  color.  Abdomen  shows  no  fluid  or  adhesions. 
Liver  slightly  decreased  in  bulk,  smooth  surface  and  sharp  edges,  hard, 
and  rusty  brown.  Gall-bladder  distended,  contained  green  fluid.  Spleen 
firm  and  normal  in  size.  Capsule  smooth,  shape  normal.  Section  sur- 
face, dark  reddish  brown,  trabeculae  distinctly  visible.  Kidney,  normal 
in  shape,  capsule  smooth,  strips  easily,  smooth  surface,  glistening,  pink- 
ish gray,  consistency  hard.  Section  surface,  poor  demarkation  between 
cortex  and  medulla.  R.  Adrenal,  thick  orange  yellow  cortex,  solid 
small  brown  medulla.  Mouth  and  teeth  normal.  Stomach  distended, 
contains  gas  and  small  quantity  yellowish  mucus.  Mucosa  everywhere 
normal.  Postmortem  blood  examined,  stained  by  Romanowsky,  but  red 
cells  were  disintegrated  possibly  by  laking  so  examination  is  not  satis- 
factoiy.  Histological  Sections :  Heart  shows  normal  epicardium  quite 
free  of  fat.  Myocardium  peculiar  in  that  fibres  immediately  under  epi- 
cardium show  marked  broadening  in  a  very  narrow  rather  sharply 
indicated  zone  where  nuclei  are  extremely  large  although  not  especially 
chromatic.  Transverse  markings  here  easily,  although  faintly  recognized, 
have  very  indefinite  borders,  their  longitudinal  fibrillas  being  ranged  in 
form  of  a  coarse  reticulum.  This  comes  about  from  frequent  and  exten- 
sive lateral  anastomoses  with  fellow  fibres  giving  appearance  of  a 
syncytium.  In  deeper  parts,  fibres  are  of  more  normal  size  but  nuclei 
are  still  large  and  fibres,  now  cut  in  tranverse  section  do  not  appear  to 
anastomose  so  freely;  there  appears  to  be  a  slight  excess  of  fibrous 


DISEASES  OF  BLOOD  AND  BONE  MARROW     101 

tissue  in  their  deeper  parts.  Arteries  quite  nonnal.  Atrophy 
with  regeneration. 

Liver. — Capsule  and  interstitial  parts  on  whole  normal.  Perilob- 
ular fibrous  tissues  largely  missing,  but  where  remaining  show  an  over- 
growth occurring  in  peculiar  zonal  arrangement  and  of  old  adult  almost 
hyaline  type.  Its  fibres  are  often  an-anged  strikingly  in  whorls.  Bile 
ducts,  ai-teries  and  veins  quite  normal.  Parenchymal  cells  of  normal 
size,  finely  granular,  prominent  normal  nuclei  and  contain  small 
quantities  of  finely  granular,  golden  brown  pigment  not  really  as 
abundant  as  commonly  seen  in  severe  anemias.  Blood  capillaries 
narrow,  contain  small  quantities  R.B.C.,  and  Kupffer's  cells  very  fre- 
quently contain  fine  granules  like  those  of  parenchymal  cells  but  of  a 
greener  tint.  In  a  few  isolated  areas  parenchymal  cells  distinctly  more 
swollen  than  others  and  many  show  disintegration  of  nucleus.  Hemosid- 
erin pigmentation.     Perilobular  fibrosis.     Slight  local  cloudy  swelling. 

Kidney. — Capsule  smooth,  interstitial  fibrous  tissue  of  organ  proper 
highly  fibrosed  in  peripheral  parts,  but  slightly  in  deeper.  No 
lymphocytic  infiltrations  anywhere  or  sclerosis  of  vessels.  Tubular 
epithelium  highly  atrophic  in  subcapsular  regions  where  tubules  are 
narrow.  In  deeper  parts  epithelium  is  at  times  so  swollen  as  to  occlude 
lumina,  where  they  are  coarsely  granular  and  occasionally  show  some 
karyolysis,  a  pink  hyaline  or  finely  granular  material.  Tufts  never 
show  fibrosis,  normal  size.  Bowman's  capsule  heavily  tliickened.  Chronic 
diffuse  nephritis   (subcapsular  type). 

Spleen. — Slightly  hyalinized  capsule,  normal  thickness.  General 
reticulum  of  pulp  slightly  fibrosed  and  poor  in  lymphocytes.  Sinuses 
broad,  crowded  with  red  blood  cells,  but  only  small  numbers  of  lympho- 
cytes. Coarse  granular  blood  pigment  abundant,  showing  greenish 
cast  on  focusing.  Maljiighian  follicles  normal  size,  slightly  fibrosed, 
and  in  several  instances  show  a  deposit  of  smooth  pink  material 
between  cells.  Congestion.  Fibrillar  fibrosis.  Hemosiderin  pigmenta- 
tion.    Local  amyloid  infiltration. 

Adrenal. — Organ  appears  nonnal  in  all  respects  save  for  presence 
of  a  few  small  irregular  areas  of  calcification  in  medulla.  These  occur 
apart  from  any  recognizable  necrotic  or  fibrous  areas.  In  one  place  one 
appears  to  lie  within  lumen  of  blood  vessel.  No  fibroses  or  special 
congestions  anywhere  in  organ,  and  cells  show  normal  details  and 
normal  numbers  of  vacuoles.    Calcareous  infiltration  of  medulla. 

Stomach. — Muscular  tunic  normal.  Submucosa  thin,  has  densely 
arranged  bundles  of  smooth,  pink  character.  Mucosa  distinctly  thinned, 
shows  comparatively  few  regions  holding  acid  cells,  consisting  for  most 
part  of  peptic  type  of  gland.  These  are  short  and  of  broader  calibre 
towards  lumen  than  deeper,  suggesting  a  hyperplasia  of  luminal  por- 
tions; stroma  richly  infiltrated  with  lymphocytes,  not  fibrosed  or  con- 
gested. Epithelium  of  crypts  has  rarefied  appearance,  shows  no  special 
degenerative  changes.     Chronic  atrophic  gastritis. 


102  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Bone  marrow  appears  as  widely  separated  large,  fat  globules  with 
intervening  granular  edematous  material  and  no  hematopoietic  ele- 
ments.   Blood  capillaries  numerous  and  highly  congested. 

Although  the  notes  fail  to  discuss  the  gross  appear- 
ance of  the  bone  marrow,  the  amount  of  alteration  in  its 
microscopy  and  the  relatively  small  output  of  pigment 
in  the  liver,  seem  to  substantiate  the  determination  of 
aplastic  anemia;  it  is  unfortunate  that  the  blood  smears 
could  not  be  used  in  the  decision.  At  all  events  the  con- 
dition of  the  intestinal  tract,  of  the  heart,  liver,  spleen, 
and  adrenal,  justify  us  in  classing  the  case  as  one  of 
primary  anemia.  The  next  and  last  instance  is  of  the 
same  type,  although  I  am  inclined  now  to  differ  from  the 
diagnoses  made  at  the  autopsy  table,  that  of  aplastic 
anemia,  and  to  place  it  in  the  hemolytic  variety.  The 
rapidity  of  the  fatal  attack,  the  redness  of  the  marrow, 
the  excessive  pigmentation,  and  the  prominence  of  recent 
degenerative  lesions  in  the  organs  are  much  more  like  the 
changes  of  a  primary  hemolytic  intoxication  than  of  an 
aplastic  anemia. 

Gray  Fox  (Canis  cinereo).  6  Weight  four  pounds,  adult,  was  in  the 
exhibition  two  years,  but  in  good  condition  until  two  weeks  before 
death,  when  it  stopped  eating  and  rapidly  fell  away. 

Diagnosis. — (Aplastic)  Primary  anemia.  Zenker's  Hyaline  of  heart 
and  skeletal  muscles.  Mucoid  degeneration  of  bone  marrow.  Conges- 
tion of  bone  marrow.  Atrophy  of  hemopoietic  elements  in  bone  marrow. 
Hemosiderin  pigmentation  of  liver.  Atrophy  of  liver.  Congestion  of 
liver.  Congestion  and  fatty  infiltration  of  kidney.  Patulous  lumina 
in  adrenal  and  absence  of  pars  glomerulosa. 

External  appearance  of  coat  good.  Decomposition  advanced  in  intes- 
tines. Skin  and  subcutaneous  tissue  faintly  yellow.  Poorly  developed, 
dark  muscles  and  fat.  Respiratory  tract  normal  throughout.  Pericar- 
dium glistening,  transparent,  and  pale,  with  no  adhesions.  Heart  a  little 
too  pale,  consistency  slightly  soft.  Normal  or  slightly  increased  size 
of  liver,  with  smooth  surface  and  sharp  edges,  consistency  friable, 
and  of  a  brownish  red  Avith  rusty  coloring.  Spleen  normal.  Kidney 
normal  in  size,  shape,  location,  and  consistency.  Capsule  strips  easily, 
and  of  a  faintly  yellow,  under  general  red,  coloring.  Adrenal  normal. 
Mouth  and  teeth  normal.  Stomach,  serosa  and  wall  normal.  Mucosa 
shiny,  autolytic,  muddy  red.  Ileum,  agminated  follicles  swollen.  Feces 
from  colon  examined  microscopically.    Pancreas  normal.    Bone  marrow 


DISEASES  OF  BLOOD  AND  BONE  MARROW     103 

tibia  and  femur  gelatinous  and  red,  not  slightest  trace  of  yellow.  Blood 
films  from  heart's  blood  show  poikiloeytosis  and  anisocytosis ;  only  one 
nucleated  red. 

Microscopic  Notes. — Heart  has  torn  but  normal  pericardium.  No 
abnormalities  of  interstitial  tissue  or  vessels.  Fibres  of  normal  width 
but  show  transverse  markings  irregularly  since  cytoplasm  becomes 
hyaline  and  swollen  in  many  places  along  its  course.  Nuclei  prominent, 
slightly  pyknotie.     Zenker's  Hyaline. 

Bone  marrow  consists  of  a  matrix  of  granular  or  fibrillar  mucoid  tis- 
sue within  the  delicate  reticulum  of  which  highly  developed  capillaries  are 
placed,  together  with  stellate  spindle  and  sealring  cells.  In  a  few 
places  only  are  myelocytes  recognizable  and  then  in  decreased  numbers. 
Plasma  cells  sometimes  found  containing  much  blood  pigment. 

Liver. — Capsule  noi-mal;  perilobular  fibrous  tissue  only  slightly 
overgrown,  moderately  infiltrated  with  lymphocytes  and  heavily  with 
blood  pigment.  Arteries,  ducts,  veins,  normal.  Parenchymal  cell  a 
little  smaller  than  normal  with  nuclei  of  normal  type,  and  crowded  with 
fine  granules  of  blood  pigment.  Latter  lie  in  usual  pericanalicular  posi- 
tion. Blood  capillaries  narrow,  moderately  congested  and  Kupffer's  cells 
also  contain  abundant  pigment  granules. 

Thyroid. — Interstitial  framework  shows  no  fibrosis  or  cellular  infil- 
trates. Blood  vessels  normal.  Acini  fairly  uniform  in  size,  none  ever 
attaining  large  proportions,  but  some  being  distinctly  below  normal. 
They  are  uniformly  filled  with  a  very  pale  pink  hyaline  material  which 
in  some  way  gradually  increases  in  color  intensity  toward  one  side, 
attaining  in  a  few  examples  usual  intensity  of  colloid.  Lining  epithelium 
is  low  cuboidal,  shows  no  special  hyperplastic  features  or  atrophy. 

Kidney. — Capsule  nonnal.  Interstitial  tissue  nonnal.  Blood  vessels 
slightly  congested.  Tubular  epithelium  granular,  disintegrated  and  fre- 
quently contains  numerous  fat  globules  and  obscured  nuclei.  Lumina 
of  about  normal  size  containing  variable  quantities  of  pink  granular 
detritus.  Glomerular  tufts  normal  in  size  and  appearance.  Subcapsular 
space  and  Bowman's  capsule  normal. 

Adrenal. — Capsule  and  pericapsular  tissue  normal.  Parenchymal 
cells  throughout  poor  or  practically  free  of  vacuoles,  such  appearing 
in  only  limited  portion  of  pars  vesicularis.  Interstitial  framework 
and  vessels  normal.  Structure  of  columns  in  pars  vesicularis  is  peculiar 
in  that  they  extend  quite  to  capsule  with  no  intervening  pars  glomerulosa, 
and  again  in  that  most  peripheral  parts  are  expanded  at  times  showing 
a  lumen,  while  deeper  parts  show  broad  cells  extending  fully  across  the 
column.  Pars  reticularis  contains  no  pigment  and  medullary  cells 
quite  normal.  Skeletal  muscles  show  comparatively  few  fibres  with 
transverse  markings.  Most  are  swollen,  hyaline,  lumpy,  and  have 
pyknotie  nuclei.     Interstitial  parts  show  no  inflammatory  change. 

Tissues  treated  by  Prussian  blue  test  for  iron.  Kidney,  adrenal,  heart 
found  to  contain  none.  Spleen,  liver  contain  abundance.  That  in  spleen 
responds  to  test  showing  that  it  is  all  iron  containing.     Two  kinds  of 


104  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

pigment  found  in  liver.  In  periphery  of  lobule  as  much  contains  iron 
as  that  which  does  not,  while  in  deeper  parts  iron  predominates.  Many 
times  both  kinds  are  recognized  in  one  cell.  On  the  whole  it  is  the  finest 
granules  which  contain  more  iron  (are  bluer)  while  iron  free  pigment 
occurs  in  bile  canaliculae.     That  in  KupfPer's  cells  stains  strongly  blue. 

Summary  of  Peimaey  Anemias. 

A  review  of  these  instances  of  grave  anemia  brings 
one  to  the  conclusion  that  there  is  a  strong  similarity  to 
the  disease  in  man.  Perhaps  we  have  constructed  a  pic- 
ture that  is  too  narrow  for  the  animal  kingdom  in  general, 
but  surely  these  few  instances  deserve  to  be  distinguished 
from  the  secondary  cases  already  presented  if  for  no 
other  reason  than  that  no  associated  etiological  condition 
was  exposed.  It  was  hoped  in  studying  the  anemias  of 
lower  animals,  and  this  hope  extends  over  all  the  subject 
of  this  book,  to  be  able  to  throw  some  light  upon  causation. 
The  thought  of  incorrect  diet  came  at  once,  but  we 
are  confronted  with  the  paucity  of  cases  among  our 
records.  Moreover,  secondary  anemia  from  digestive 
and  dietetic  troubles  is  clear,  but  how  we  can  use  this 
argument  for  an  essential  change  in  hematopoiesis  and 
natural  hematolysis,  is  far  from  evident.  It  will  be 
noticed  that  I  have  studiously  avoided  grouping  any  case 
with  parasites  among  the  primary  cases,  nor  will  there 
be  found  any  evidence  of  generalized  infectious  disease. 
Most  of  the  reported  instances  of  pernicious  anemia  in 
the  lower  animals  have  been  associated  with  one  or  other 
of  these  factors,  although  certain  authors  (Kitt,  Hutyra 
and  Marek)  maintain  that  a  causeless  variety  prob- 
ably exists. 

Leucemia. 

This  condition  is  fairly  well  recognized  by  veter- 
inarians as  occurring  among  domesticated  animals,  but 
in  the  records  of  this  Garden  it  has  occurred  rarely, 
indeed  only  once  in  a  mammal  and  butj  five  times  in  birds. 
It  is  interesting  that,  in  the  wealth  of  material  at  the  dis- 


DISEASES  OF  BLOOD  AND  BONE  MARROW     105 

posal  of  Plimmer  and  his  associates,  only  one  case,  a  pole- 
cat with  lymphatic  leucemia,  is  noted,  and  but  very  few 
avian  instances.  Herewith  is  submitted  the  protocol  of 
our  single  mammalian  case;  perhaps  we  have  missed 
others  of  a  mild  grade  dying  during  the  early  stages 
because  their  resistance  to  infection  was  reduced. 
Unfortunately,  perhaps  because  of  the  postmortem 
changes,  but  more  likely  because  it  appeared  at  first  as  if 
we  had  to  do  with  a  case  of  generalized  tuberculosis, 
the  bone  marrow  in  this  case  was  not  examined.  Never- 
theless the  infiltrative  character  of  the  lesions,  the  absence 
of  distinct  tumors  and  the  numerous  mononuclears  in  the 
blood  as  seen  in  sections  seem  to  justify  a  diagnosis  of 
leucemia,  in  all  probability  of  the  lymphatic  type.  There 
follows  this  case  one  with  similar  gross  and  microscopic 
picture  which  has  no  visible  increase  of  leucocytes,  but  a 
very  distinct  myeloid  picture  in  many  places. 

Common  Opossum  {Didelphys  virginianus).  6  Adult.  No 
evidence  of  illness.  Found  dead.  Lymphatic  leucemia  (involv- 
ing all  viscera  and  lymph  nodes).  Diffuse  nephritis.  Both  lungs  have 
become  entirely  involved  in  a  firm,  gray  yellow  mass  not  adherent  to 
any  serous  surface.  Practically  no  normal  lung  tissue  is  left.  This 
seems  like  tuberculosis  but  no  tubercle  bacilli  could  be  found  in  a  good 
smear.  Estimation  of  the  normal  cubic  capacity  of  an  opossum  lung 
was  made  to  be  about  five  cubic  inches.  In  this  case  not  over  one-half 
cubic  inch  remained  respirable.  Bronchial  lymphatic  glands  were  en- 
larged, firm,  yellow  gray,  with  no  recognizable  lymphatic  tissue.  Heart 
muscle  was  firm  and  flaccid,  pale  and  striated.  Liver  very  large,  firm 
and  tough,  with  smooth  surface  and  sharp  edges.  Color  pale  brown. 
Section  surface  glistening,  dry,  smooth,  opaque.  Common  bile  duct 
patulous.  Spleen,  slightly  increased,  firm,  tough  consistency,  capsule 
smooth.  Section  surface,  smooth,  firm,  brown-red,  pale  pulp,  prominent 
follicles,  and  trabeculse  faintly  visible.  Right  kidney,  slightly  decreased, 
normal  in  shape.  Capsule  smooth,  strips  with  difficulty,  tears  surface. 
Surface,  granular,  color  brown,  consistency  firm.  Thickness  of  cortex, 
narrow  irregular,  markings  irregular  and  obscure.  Small  mass  of 
fibrous  material  in  cortex  about  3x3  mm.  like  those  in  lungs.  There 
are  also  numerous  pale  yellow-gray  areas  in  cortex  and  outer  medulla, 
round  and  streaky,  distorting  the  striated  architecture.  Right  adrenal 
converted  to  a  yellow  gray  mass  like  lungs.  The  mesenteric  and  retro- 
peritoneal lymph  nodes  are  firm,  gray-yellow.  This  includes  those  under 
diaphragm  and  around  coeliac  axis. 


106   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

The  histology  of  the  organs  may  be  described  together.  The  infil- 
trate described  is  a  densely  packed  mass  of  large  cells  with  large,  well 
staining  nuclei  and  a  very  narroAv  rim  of  protoplasm.  It  is  not  limited 
by  any  definite  wall  or  septa.  It  has  no  interstitial  tissue.  There  is  no 
blood  supply  in  the  densest  masses  but  the  walls  of  the  blood  vessels 
remain  intact  Avherever  the  mass  surrounds  them.  In  the  lungs  it  has 
involved  all  structures  indiscriminately,  and  has  destroyed  practically 
all  of  the  respiratory  surface.  It  seems  to  follow  by  preference  the 
peribronchial  space.  A  few  glands  may  be  seen  in  the  centre  of  this 
mass,  but  they  are  rapidly  undergoing  degeneration.  In  the  liver  the 
infiltrate  is  chiefly  beneath  the  capsule  extending  inward  but  a  very 
short  distance.  There  are  no  large  masses  as  in  other  organs  but  small 
infiltrates  are  seen  at  the  portal  areas.  The  spleen  shows  a  diffuse 
excess  of  pulp  cells  and  many  of  the  cells  above  described,  the  difference 
being  only  in  the  size  of  the  nucleus  which  is  smaller  in  the  pulp  cells. 
There  are  very  small  round  cells  relatively.  Follicles  are  absent,  connec- 
tive tissue  not  altered.  Note  states  follicles  prominent;  this  is  due  to 
nodal  hyperplasia  of  the  large  mononuclears  above  described.  The 
parenchyma  in  the  kidney  is  anemic,  the  epithelium  is  slightly  pigmented 
but  this  is  probably  not  abnormal.  Between  the  tubules  especially  of 
the  outer  layer  of  the  medulla  and  medullary  ray  but  also  in  the  cortex 
and  around  the  glomeruli  are  diffuse,  irregular,  infiltrating  masses  of 
the  cells  as  described  above.  In  some  places  in  the  kidney  hyaline  casts 
are  being  formed  probably  due  to  the  degeneration  of  the  epithelium 
by  pressure.  There  are  a  few  distentions  of  the  tubules.  The  capsular 
space  is  free.  In  the  neighborhood  of  the  collections  the  capsules  are 
a  trifle  thicker  than  normal.  Lymph  nodes,  similar  to  spleen  in  that 
most  of  the  bodies  are  thoroughly  oveiTun  with  the  large  mononuclears. 
The  sinuses,  both  marginal  and  internal,  are  practically  obliterated  by 
these  cells.  In  the  blood  vessels  of  the  lungs  and  liver  there  are  many 
large  mononuclears,  perhaps  not  as  large  a  number  as  might  be  seen 
in  leucemia,  but  decidedly  in  excess  of  normal. 

Common  Marmoset  {Callithrix  jacchiis).  6  Adult.  Had  cage 
paralysis  for  two  months  before  death  and  declined  gradually  from 
that  time. 

Diagnosis. — Bronchopneumonia.  Myeloid  hyperplasia  of  bone  mar- 
row. Myeloma  in  pancreas.  Fatty  degeneration  of  liver.  Constipation. 
Nematodes  in  cecum.  Animal  is  thin,  skin  bare  in  spots.  Both  lungs  are 
pale  pink  with  large  areas  of  deep  red  consolidation.  Heart  is  dilated, 
increased  in  size  with  firm,  red-brown  muscle.  Liver  is  firm,  red-brown, 
with  smooth  surface  and  sharp  edges.  Section  surface  is  glistening, 
smooth  and  moist.  Lobular  outlines  are  clear  by  reason  of  pale  lines. 
The  gall-bladder  is  normal  in  size  and  contains  fluid  pale  green  bile; 
duct  patulous.  Spleen  is  normal  in  size,  smooth  capsule,  soft,  purple 
pulp,  follicles  small  and  faint,  trabeculae  fairly  prominent.  Kidneys 
normal  in  size  and  shape.  Capsule  smooth.  Section  surface  smooth  and 
broAvn  and  firm.     The  glistening  section  surface  has  a  narrow  cortex, 


DISEASES  OF  BLOOD  AND  BONE  MARROW     107 

swells  slightly,  with  prominent  striae.  Intestines  throughout  are  pale 
on  serosa.  Wall  thin.  Mucosa  flat,  pale  pink.  Contents  creamy  mucus 
in  the  upper  intestine.  Large  intestine  contains  large  masses  of  very 
firm  feces.  Cecum  is  distended  with  feces  and  a  great  quantity  of 
nematode  worms.  They  are  not  attached  to  mucosa  nor  does  mucosa 
seem  altered  because  of  their  presence.  Skeleton  and  muscles. — Long 
bones  of  extremities  break  easily,  but  Avith  snap.  Skull  can  be  dented 
with  fingers.     Bone  marrow  of  femur  bright  red. 

Microscopical  Notes. — Liver  shows  moderate  degree  of  fatty  degen- 
eration with  capillary  congestion.  Kidneys  negative.  Some  postmortem 
change  in  last  two  organs.  Spleen,  marked  congestion.  Hyperplasia  of 
large  lymph  cell  type,  particularly  in  follicular  centres.  Blood  destruc- 
tion moderate.  Bone  marrow  seen  in  condition  of  marked  activity  of 
myeloid  type.  Aside  from  enormous  crowding  of  strands  there  does  not 
seem  to  be  any  atypical  cell.  Intestines  show  practically  no  change. 
Same  condition  holds  in  pancreas.  In  several  places  in  pancreatic  ducts 
cross  sections  of  nematodes  may  be  found.  In  among  lobules  of  pan- 
creas is  a  well  encapsulated  cellular  mass  without  particular  architecture. 
It  consists  of  cells  of  large  lymphocyte  or  endothelioid  series.  There  are 
numerous  cells  of  size  and  staining  characters  of  small  lymphocytes. 
There  are  no  megalocytes  but  there  are  some  indistinguishable  from 
myelocytes.  This  may  be  an  intrapancreatic  lymph  node.  One  small 
lymph  node  found  in  section;  it  shows  a  picture  quite  like  the  marrow 
except  for  megalocytes.  Blood  vessels  do  not  show  an  excess  of  leuco- 
cytes in  free  or  coagulated  blood. 

Perhaps  this  latter  case  belongs  to  the  aleucemic 
leucemias  or  pseudoleucemias.  These  two  conditions  are 
recognized  by  the  difference  in  circulating  leucocytes,  a 
piece  of  information  not  at  our  disposal.  The  whole  sub- 
ject of  hemato-lymphatic  affection  must  remain  unsettled 
in  so  far  as  a  diagnostic  name  is  concerned,  for  in  very 
few  cases  has  the  blood  of  our  animals  at  autopsy  been 
in  a  state  permitting  reliable  observations  upon  stained 
smears,  because  of  coagulation,  lysis  or  decomposition. 
After  considering  a  few  more  of  the  diseases  of  the  blood 
and  marrow,  the  lymphatic  apparatus  will  be  considered. 
But  there  is  a  borderland  to  which  a  word  might  be 
devoted  at  this  time,  that  group  to  which  various  names 
— Hodgkin's  disease,  pseudoleucemia,  general  adenop- 
athy, adenie,  aleucemic  leucemia — have  been  applied 
and  which  has  been  accepted  as  occurring  in  the  domesti- 
cated animals.     Since  I  have  been  occupied  for  several 


108   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

years  in  a  study  of  this  clinico-pathological  complex  in 
the  human  being,  such  cases  have  been  searched  for  most 
diligently,  but  without  success.  The  New  York  Zoological 
Park  records  a  case  of  Hodgkin's  disease,  mthout  specifi- 
cations, in  1901,  and  at  the  London  Garden  a  pseudo- 
leucemia  was  found.  The  paucity  of  leucemia  and 
of  the  aleucemic  adenopathies  in  lower  animals  and 
their  relative  frequency  in  man  excite  speculation  as 
to  their  interdependence;  but  more  of  this  under 
the  lymphatics. 

Avian  Leucemia. 

The  class  Aves  is  rather  better  represented  in  the 
group  of  leucemias,  but  here  the  well  known  infectious 
disease  may  confuse  the  picture.  The  birds  affected  were 
Psittaci  3  (1  parrot,  1  parrakeet,  and  1  amazon),  Herodi- 
ones  (stork)  1,  and  Galli  (Gambel's  quail),  1.  There  was 
no  close  association  of  these  cases  either  in  time  or  hous- 
ing. One  of  the  parrots  and  the  stork  had  a  picture 
suggesting  that  given  by  Warthin  for  avian  leucemia 
while  the  remainder  presented  greater  evidence  of  a 
generalized  infection,  such  as  Moore  described,  associated 
with  the  finding  of  the  B.  sanguinarium ;  this  organism 
was  isolated  once,  but  no  secondary  cases  succeeded  upon 
the  death  of  this  bird.  It  seems  hardly  profitable  to  quote 
protocols  of  this  relatively  unimportant  condition,  espe- 
cially since  it  is  fairly  well  known. 

The  separation  of  the  two  groups  just  specified  might 
be  discussed,  however,  for  it  is  by  no  means  certain  that 
they  are  or  are  not  different.  When  a  pathological 
picture  of  leucemia  gives  a  decided  impression  of  an 
acute  infection  there  are  very  prominent  involvements 
of  the  viscera  but  no  lymph  nodal  masses.  On  the  other 
hand,  in  the  cases  mth  nodular  masses  corresponding  to 
( the  scanty  lymph  tissue  of  birds,  there  is  much  less  infil- 
trative involvement  of  viscera  and  less  parenchymatous 
degeneration.     This    suggests   that   they   are   different 


DISEASES  OF  BLOOD  AND  BONE  MARROW     109 

processes,  but  an  analogous  contrast  may  be  found  in  the 
pathologic  anatomy  of  acute  and  chronic  leucemia  in  man, 
and  I  am  inclined  to  view  them  as  stages  of  the  same  dis- 
ease. In  one  of  our  infectious  cases  noted  above  the  lesion 
was  certainly  myelogenic  for  the  infiltrate  in  the  organs 
and  the  cells  in  blood  smears  showed  an  enormous  number 
of  eosinophilic  and  basophilic  polynuclears  greatly  in 
excess  of  normal.  The  study  of  two  of  our  cases  confirms 
the  picture  as  given  for  lymphatic  and  myeloic  leucosis 
by  Ellermann  (1),  but  material  corresponding  to  his 
lymphoidocytes  or  erythroleucotic  group  has  not  come  to 
our  attention.  Cells  with  deeply  staining  basophilic 
protoplasm  and  a  lymphoid  nucleus  are  certainly  to  be 
found  with  reasonable  ease  in  the  avian  marrow  normally 
and,  more  than  this,  can  be  detected  by  careful  search  in 
nearly  all  cellular  infiltrates  of  organs  not  leucemic  in 
nature.  Perhaps,  as  Ellermann  states,  they  are  collateral 
stages  in  normal  erythrogenesis. 

The  Bone  Makeow. 

Since  the  foregoing  conditions  so  vitally  concern  the 
bone  marrow,  it  is  but  natural  to  give  to  this  structure 
a  separate  consideration.  From  what  is  known  of  the 
origin,  physiology,  anatomy  and  regeneration  of  the 
marrow  from  the  work  of  Ponfick,  of  Neusser,  Bunting, 
Selling,  Werigo  and  many  others,  it  seems  highly 
probable  that  the  principal  conclusions  reached  in  the 
study  of  human  medicine  and  experimental  pathology, 
apply  to  the  whole  group  of  animals  here  under  discus- 
sion. The  peculiar  arrangement  already  mentioned  as 
encountered  in  the  marrow  of  birds  differs  little  if  any 
from  the  eiythropoietic  centres  seen  in  man  after  experi- 
mental anemia,  although  it  may  be  somewhat  more 
orderly.  Myeloblasts  or  megakaiyocytes  are  not  numer- 
ously present  in  any  order,  but  seem  more  prominent  in 
the  mammals  than  in  birds.    In  so  far  as  the  mononuclear 

( 1 )    The  Leucoses  of  Foicls,  London,  1922. 


no   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

groups  are  concerned,  one  can  state  with  reasonable 
certainty  that  they  differ  little  throughout  the  animal 
kingdom.  They  occur  in  islands,  strands,  or  infiltrate — 
like  groups,  are  mixed  granular  and  non-granular  in 
character  and,  with  exception  of  the  frankly  oxy- 
philic cells,  are  distinctly  basic  in  tinctorial  affinity. 
In  a  case  probably  myeloma,  soon  to  be  discussed, 
there  is  not  a  single  acidophilic  or  multinucleated 
cell  to  be  found  in  two  sections.  As  might  be  expected 
from  the  greater  eosinophilic  content  of  the  circulating 
blood  in  the  Aves,  greater  numbers  of  such  cells  are  to  be 
found  in  the  marrow  and  they  are,  understandingly 
enough,  sometimes  found  in  distinct  nodes  and  groups 
containing  mono-  and  polynuclear  varieties.  Concerning 
the  platelets,  no  sufficient  data  are  at  our  disposal  to  war- 
rant a  general  statement.  In  the  avian  marrow  they  can 
be  made  out  quite  clearly,  as  in  the  blood,  and  have  a 
greater  diameter  and  a  sharper,  more  chromatic  nucleus 
than  in  the  higher  mammalian  blood.  Mast  cells  are  quite 
common  in  lower  mammals  and  birds,  in  whose  blood  they 
maintain  an  appreciable  percentage,  while  in  the  marrow 
they  stand  out  clearly.  It  is  noteworthy,  in  the  light  of 
Graham's  statement  that  the  hemic  basophile  is  but  a 
degeneration  form  of  the  eosinophile,  that  in  the  bird's 
marrow,  large  mono-  and  polynuclear  cells  ^\dth  both 
kinds  of  granules  easily  may  be  found  by  Eomanowsky 
stain.  <  An  increase  of  eosinophiles,  seen  in  avian  as  well 
as  in  human  parasitism,  is  not  necessarily  accompanied 
by  basophilia.  ~ 

Hyperplasia  and  atrophy  of  marrow  in  the  lower 
mammals  follow  much  the  same  conditions  as  in  higher 
groups.  During  acute  general  infections,  as  by  the  para- 
colon bacillus  in  carnivores,  it  is  common  to  find  a  distinct 
increase  in  the  mononuclear  centres,  while,  in  suppu- 
rative lesions  a  polynucleosis  results.  'The  bird, 
however,  responds  less  readily  with  leucocytes,  judged 
by  cross  sections  of  blood  vessels  and  the  activity  of  the 


DISEASES  OF  BLOOD  AND  BONE  MARROW     111 

marrow.  The  latter  may  show  a  myeloid  picture,  but 
mononuclears  without  granules,  with  deeply  staining 
nucleus  and  protoplasm,  are  usually  more  numerous ;  two 
cases  recently  studied,  one  of  tuberculosis  and  one  of 
pneumonia  with  general  congestion,  had  similar  bone 
marrow — pale  homogeneous  red  with  distinct  mono- 
nucleosis, more  outspoken,  however,  in  the  former  case. 
Pigment  is  not  common  in  the  avian  marrow. 

The  relation  of  the  marrow  to  general  conditions  has 
been  mentioned  in  the  foregoing  pages,  but  perhaps  the 
following  diagnoses  will  illustrate  other  connections  seen 
among  our  records : 

Bactrian  Camel  {Cameliis  hactrianus) . — Hydatid  dis- 
ease of  lung  and  liver.  Hemorrhagic  enteritis.  Atrophic 
bone  marrow.   Calcified  areas  in  thyroid. 

American  Gray  Wolf  {Canis  mexicanus) . — Cretinoid. 
Hemorrhagic  external  pachymeningitis  with  cranio- 
tabes.  Secondary'  hyperplasia  of  thyroid  with  colloid. 
Chronic  IjTuphatic  hyperplasia.  Chronic  interstitial 
nephritis.  Chronic  enteritis.  Osteogenesis  imperfecta. 
Hemorrhagic  bone  marrow.  Concentric  hypertrophy  and 
dilatation  of  heart. 

Myeloma. 

Perhaps  no  pathological  condition  has  given  rise  to 
more  varied  opinions  than  the  tumor-like  hyperplasias  of 
the  bone  marrow,  growths  resembling  bone  sarcoma  with 
and  without  giant  cells — ^myeloma,  chloroma,  pseudo- 
leucemia  ossium  and  many  others.  In  brief  only  two 
cases  occur  in  our  series  which  could  be  admitted  to  this 
category.  There  have  been  osteomata,  but  they  were  so 
clearly  local  tumors  that  they  cannot  be  included  in 
myeloid  neoplasms  that  are  assumed  to  be  systemic  in 
nature.  Here  is  not  the  place  to  engage  in  a  discussion  of 
the  correct  classification  and  nomenclature  since  there 
are  included  only  the  aleucemic  newgro^vths  usually 
assumed  to  originate  from  blood-making  cells.    The  first 


112   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

case  seems  to  be  a  myeloma  because  of  the  involvement  of 
many  bones  and  the  infiltrates  in  the  liver.  Judging  by 
the  cross  section  of  blood  vessels  there  is  no  leucemia,  but 
of  course  this  is  not  final,  although  somewhat  supported 
by  the  normal  size  of  the  spleen  and  lymph  nodes; 
lymphatic  structures  need  not  be  enlarged  in  myeloid 
leucemia  although  they  usually  are. 

Beechy's  Gopher  {Citellus  gramtnurus  beecheyi).  Adult  6. 
Gradual  loss  of  power  in  limbs  beginning  about  two  weeks  ago.  No 
other  symptoms. 

Diagnosis. — Multiple  myeloma.     Anemia.     Acute  parenchymatous 
nephritis.     Chronic  ulcerative  gastritis. 

External  Appearance. — General  condition  fair,  hair  in  good 
condition.     Fully  developed  animal.     Muscles  contained  no  fat. 

Respiratory  Tract. — Thymus,  large  soft  gray  homogeneous.  Both 
lungs  are  distended,  pale,  homogeneous,  yellow  pink,  boggy,  do  not 
crepitate,  but  contain  no  edema.  Lymph  nodes,  small  soft  anthraeotie. 
Pericardium  contained  no  fluid. 

Abdomen. — No  adhesions.  Size  of  liver  normal  or  slightly  small, 
ifii-m  and  pale  brown-red.  Architecture  irregular  showing  areas  of 
perfect  homogeneity  and  others  where  lobules  are  clear  outlined  by 
paler  interstices.  Gall-bladder  distended,  contains  brown  fluid.  Com- 
mon bile  duct,  patulous.  Spleen,  normal  or  slightly  small,  consistency 
soft,  capsule  smooth,  location  normal;  section  surface,  homogeneous 
pale  pulp,  faint  trabeculse.  Kidney  normal  in  size  and  shape,  capsule 
smooth,  strips  easily,  smooth  brown  surface ;  section  surface  glistening, 
opaque;  consistency  firm;  thickness  of  cortex  normal  and  of  medulla 
normal;  homogeneous  cortex,  glomeruli  not  visible. 

Adrenal. — Narrow,  brown,  opaque  cortex;  pale  brown,  opaque 
medulla.  Bladder,  small  quantity  of  cloudy  urine  showing  albumin, 
bile  and  many  granular  casts,  few  hyaline  casts.  Teeth  carious  broken ; 
mouth  pale.  Stomach  distended;  serosa  and  wall  pale,  contained  gas; 
mucosa  pale,  flat,  translucent  except  in  pyloric  segment  where  it  is 
slightly  thickened,  irregular,  opaque  and  there  are  several  saucer  shaped 
depressions  covered  with  black,  shiny  material ;  these  seem  like  sluggish 
ulcers.  Intestinal  tract  throughout  seems  normal  save  for  its  pallor. 
Few  natural  fecal  masses  in  lower  gut. 

Skeleton  and  Muscles. — All  bones  are  irregular  in  thickness,  very 
brittle  and  show  in  their  length  irregular  swellings  made  up  of  perios- 
teal groAvth  and  probably  increase  in  marrow.  The  latter  is  pale  yellow 
with  punctate  homon-hages.  Skeleton  seems  too  soft  to  give  support, 
but  there  is  little  deformity. 

Microscopy. — Bone  section  shows  a  myelomatous  growth  of  costal 
marrow,  new  myelocytes  predominating.  The  cells  are  packed  in  dis- 
orderly fashion  through  the  marrow.     They  are  chiefly  lightly  granular 


DISEASES  OF  BLOOD  AND  BONE  MARROW     113 

but  a  few  distinct  promyelocytes  are  found.  The  hyperplasia  is  invading 
cartilaginous  bone  with  absorption.  In  some  places  cartilage 
is  of  fetal  type.  Perichondrium  is  active  but  there  is  no  round  cell 
infiltration.  Ossification  is  imperfect  at  costochondral  junctions.  Kidney 
shows  granular  and  vacuolar  degeneration  of  epithelium  with  flattening 
of  tubular  lining.  Nuclei  are  for  most  part  normal.  Epithelium  of 
tufts  and  Bowman's  capsule  is  likewise  granular  and  vacuolated.  Dis- 
tal and  discharging  tubules  seem  to  be  most  affected.  No  well  defined 
casts  are  found. 

Liver. — The  fine  markings  mentioned  in  the  notes  con-espond  to 
areas  of  infiltration  of  large  pale  mononuclears  without  granules.  There 
is  no  fibroblastic  or  polynuclear  increase  around  them.  Very  small 
necroses  seen  in  hepatic  lobules.  Very  slight  increase  of  connective 
tissue  is  noted. 

This  seems  to  be  a  tumor  of  true  myelocytic  origin; 
none  of  the  cells  was  of  the  plasma  type  as  usually 
depicted.  The  second  case  stands  in  the  files  as  a  mye- 
loma, yet  the  full  description  and  slides  have  been  mislaid. 
It  is  cited  briefly  for  record. 

Samoli  Ostrich  {Struthio  moJybdophanes) .  Adult  6.  Would  not 
eat  for  three  weeks. 

Diagnosis. — Tuberculosis  of  lung,  liver,  spleen,  kidney,  mesenteric 
and  cervical  lymph  glands.    Myeloma  of  periosteum  of  pelvis. 

Skeleton  and  Muscles. — Large  tumor  mass  lying  on  inside  of  pelvis 
measuring  about  ten  inches  by  twelve  inches  with  a  thickness  of  about 
four  inches.  It  is  sharply  defined  and  separated  from  adjacent  muscle 
by  a  capsule.  Inner  border  is  directly  below  the  peritoneum,  and  outer 
border  lies  directly  on  bony  pelvis.  Traced  to  its  origin  it  seems  to 
come  from  pelvis  yet  tumor  peels  off  bone  easily,  leaving  a  rough  sur- 
face. The  tumor  is  soft  and  succulent,  of  a  red-gray  color  and  contains 
some  fat  and  much  irregularly  placed  masses  of  bony  tissue. 


SECTION  V 

DISEASES  OF  THE  LYMPHATIC  TISSUES 
INCLUDING  SPLEEN 

Since  we  have  followed  diseases  of  the  blood  from  the 
simple  anemias  to  a  place  where  mononucleosis  in  the 
circulatory  system  and  in  the  fixed  tissues  is  the  promi- 
nent feature,  another  step  reaches  the  area  whence  most 
of  these  cells  emanate — the  lymphatic  system.  The 
anatomical  and  physiological  position  of  the  lymphatic 
circulatory  apparatus  is  closely  analogous  in  the  classes 
under  discussion,  and  it  stands  in  an  anatomico-clinical 
sense,  closer  to  the  hematopoietic  system  than  to  any 
other  structures,  throughout  the  whole  animal  creation. 
This  anatomical  division  of  the  circulation  is  closely  com- 
parable, for  pathological  purposes,  in  the  mammals 
whereas  in  the  birds  one  finds  noteworthy  variations. 
In  the  class  Aves  lymphatic  radicals  are  extremely 
numerous,  the  plexuses  in  the  extremities  and  thorax 
perhaps  being  complemental  for  the  rather  scantily  sup- 
plied blood  vessels.  About  the  fibulo-tibial  and  femoral 
muscles  the  tiny  lymph  vessels  form  an  extraordinarily 
dense  and  intricate  lacework,  a  replica  of  which  may  be 
found  in  muscles  of  the  upper  limb,  while  in  the  pelvis 
and  thorax  a  rich  plexus  is  distributed  around  both  kinds 
of  blood  vessels  and  also  lies  within  the  walls  of  air  sacs. 
The  air  sac  walls  in  the  chest  display  lymphatic  lines 
very  well  in  birds  that  have  been  for  many  years  exposed 
to  railway  dust,  the  natural  pearl  gray  glistening  mem- 
brane looking  as  if  black  pepper  had  been  evenly  dusted 
over  it.  The  lymph  glands  or  compound  nodes  so  easily 
discovered  in  mammals  are  practically  absent  in  birds. 
Along  the  large  thoracic  vessels  and  in  the  pelvis  of  some 
anserine  and  struthious  varieties,  small  illy  ■  defined 
masses  of  lymphatic  tissue  may  be  discovered  by  careful 

114 


DISEASES  OF  THE  LYMPHATIC  TISSUES     115 

search  but  they  do  not  possess  nodal  arrangement  and 
capsule.  Lymph  folhcles  on  the  other  hand,  are 
quite  numerous  in  the  respiratory,  and  especially,  the 
intestinal  tract.  Groups  of  follicles  may  also  be  found 
in  the  lining  of  the  upper  ends  of  these  tracts  in  such 
situations  that  the  names  faucial,  pharyngeal,  or  even 
tubal  tonsils  are  justly  applied.  I  do  not  find  any  ref- 
erence to  a  lingual  tonsil  in  birds,  but  this  structure  is 
found,  with  of  course  many  modifications,  do\\Ti  as  far  as 
the  monotremes.  The  lymphoid  tissue  of  the  naso- 
pharyngeal region  is  so  placed  in  animals  as  to  be 
exposed  to  aerial  and  food  infections,  just  as  it  is  in  man. 
It  is,  however,  noteworthy  that  chronic  inflammations 
leading  to  hypertrophy  or  to  obstruction  have  not  come 
to  our  attention.  There  is  only  one  diagnosis  of  chronic 
tonsilitis  in  our  records,  a  determination  based  upon  the 
nodular  red  brown  prominence  of  the  tonsillar  region,  but 
there  were  no  true  hypertrophy  and  areas  of  fibrosis  com- 
bined with  necroses  as  seen  in  human  medicine ;  this  case 
concerned  a  Chacma  baboon. 

Hypeeplasia. 

In  so  far  as  the  reaction  to  infectious  or  toxic  agents 
is  concerned  there  seems  to  be  a  fairly  uniform  character 
through  the  mammalian  groups,  but  in'  the  Aves  there 
are  a  few  differences  worthy  of  mention.  It  is  at  once 
admitted  that  these  observations  upon  birds  are  based 
upon  a  very  few  sections  of  isolated  lymphatic  tissue, 
but  they  are  supported  by  records  of  the  changes  in  nodes 
in  mucous  membranes  and  viscera,  and  by  those  in  the 
spleen.  The  first  and  perhaps  most  noteworthy  difference 
is  in  the  paucity  in  birds  of  large  mononuclears  of  the 
endothelial  type.  Their  position  in  chords,  sinuses  and 
germinal  centres  is  taken  by  deeply  staining  mono- 
nuclears, of  the  size  and  general  character  of  large 
hnnphocytes  as  seen  in  the  blood.  The  hyperplasia  in  the 
follicles  is  much  more  dense  but  it  is  outdone  by  that  in 


116   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

the  chords.  Since  the  nodal  tissue  of  birds  is  not  so 
sharply  delimited  by  some  sort  of  capsule,  it  is  but 
natural  that  the  hyperplasia  should  be  diffuse;  in  the 
intestinal  wall  it  may  extend  laterally  twice  or  thrice  the 
width  of  the  normal  follicle.  Necrosis,  unless  the  disease 
be  mycotic,  tuberculous  or  parasitic,  is  uncommon. 

It  may  be  well  to  discuss  for  a  space  the  reaction  to 
infection  of  the  thoracic  and  intestinal  lymphatics  in 
mammals.  The  amount  of  lymphadenoid  tissue  in  the 
mediastinum  is  very  great  in  some  mammals,  notably  the 
Ungulates,  while  in  others,  the  Eodents  and  Primates,  for 
example,  it  is  not  so  plentiful.  Nevertheless  the  gross 
and  minute  changes  are  usually  of  the  edemato-exudative 
type — large,  piiik,  soft,  moist  glands.  In  the  abdomen, 
on  the  other  hand,  one  usually  finds  well  outlined,  firm, 
yellow  nodes  in  the  mesentery  and  behind  the  peritoneum. 
This  is  not  only  to  be  discovered  in  various  chronic 
diseases  of  the  intestine  but  even  in  acute,  so-called  toxic 
enteritis  seen  in  carnivores  from  food  poisonings.  One 
must  therefore  ask  if  the  local  nodes  abundant  in  the 
intestinal  mucosa  do  not  take  up  the  poisons  which  cause 
the  acute  hyperplasia  and  are  in  turn  backed  up  by  the 
stalk  glands.  Even  in  so  acute  and  overwhelming  a 
disease  of  the  intestine  as  hog  cholera  (which  we  have  not 
had  here)  the  glands  retain  their  general  structure, 
although  hemorrhagic,  while  in  late  stages  they  become 
firm,  sharply  outlined  and  pigmented.  In  birds  there  may 
be  swelling  of  tlie  omental  bursas,  but  as  there  are  no 
IjTnph  glands  no  masses  are  found.  ' 

The  response  on  the  part  of  the  lymphatic  tissues  as  a 
system,  or  some  large  section  of  this  system,  is  shoAvn  in 
the  following  figures.  It  is  our  practice  to  include  in  the 
diagnosis  general  acute  or  chronic  lymphadenitis  or 
inflammation  of  a  large  drainage  area.  Acute  changes 
have  been  mentioned  103  times,  in  which  the  important 
orders  are  represented  as  follows:  Primates  21,  Car- 
nivora  46,   Ungulata   15,   Rodentia   3,   Marsupiaha   13, 


DISEASES  OF  THE  LYMPHATIC  TISSUES     117 

Pinnepedia  4.  Chronic  changes  are  mentioned  43  times 
as  follows :  Primates  7,  Carnivora  19,  Ungulata  14,  Mar- 
supialia  1,  and  Pinnepedia  2.  This  great  proportion 
among  the  Carnivora  does  not  indicate  that  they  have 
more  lymphatic  structures  for  such  an  advantage  is  prob- 
ably possessed  by  the  Ungulata,  but  perhaps  should  be 
interpreted  as  an  evidence  for  this  order  of  the  ready 
response  to  irritation  on  the  part  of  the  tissues  in  ques- 
tion. They  probably  suffer  more,  as  we  shall  see  later,  with 
inflammation  affecting  drainage  tracts.  The  hyperplasias 
or  inflammations  included  in  the  figures  above  most  often 
accompany  gastroenteritis,  pulmonary  diseases  or  long 
standing  infectious  processes  such  as  arthritis,  while 
there  are  also  lymphatic  enlargements  both  local  and 
general,  associated  with  skeletal  degenerations  (rachitis- 
osteomalacia)  and  with  thyroid  disease.  The  former  may 
be  described  as  lymphadenitis,  the  latter  as  lym- 
phatic hyperplasia. 

Unlike  lymphadenitis,  a  condition  associated  with 
some  definite  infectious  or  toxic  cause,  systemic  hyper- 
plasia of  the  lymphatic  tissue  may  be  apparently  primary 
and  causeless.  In  a  pathological  and  chnical  sense  alike 
these  hyperplasias  are  protean  in  their  manifestation, 
making  a  satisfactory  classification  extremely  difficult. 
For  our  purposes  they  are  divisible  into  acute  and 
chronic,  associated  with  an  increased  number  of  circu- 
lating lymphocytes  and  without  such  a  lymphocytosis. 

The  first,  acute  systemic  lymphatic  hyperplasia,  is 
known  in  man  as  status  thymico-lymphaticus,  a  well 
recognized  condition  chiefly  encountered  in  youthful 
males  having  some  of  the  stigmata  of  the  opposite  sex. 
There  is  no  record,  nor  have  I  any  recollection  of  a  patho- 
logical state  in  a  wild  animal  comparable  to  this  condition. 

If  acute  generalized  lymph  node  increase  be  associated 
mth  lymphocytosis,  acute  lymphatic  leucemia  exists; 
there  is  no  case  in  our  records.  Chronic  enlargement  of 
lymph  nodes  with  increase  of  circulating  mononuclears  is 


118   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

chronic  lymphatic  leucemia ;  a  case  of  this  has  been  cited 
under  leucemia. 

Lymphomatosis. 

Chronic  enlargements  of  the  lymphatic  tissues  without 
leucemic  blood  present  a  be-svildering  number  of  varieties, 
the  best  known  names  of  which  are  Hodgkin's  disease, 
aleucemic  leucemia,  pseudoleucemia  and  lymphomatosis. 
On  occasion  they  are  at  first  localized  tumors,  being 
generalized  only  late  in  the  course ;  under  these  circum- 
stances they  are  usually  classed  wdth  neoplasms  in  the 
form  of  leucosarcoma  and  lymphosarcoma.  Examples 
of  lymphatic  disease  answering  the  above  description  are 
limited  to  two,  but  even  these  must  be  explained  in  certain 
particulars.  There  has  already  been  quoted  under 
leucemia  a  case  of  a  common  Marmoset  (Callithrix  jac- 
clius)  which  was  probably  aleucemic  leucemia  of  the 
myeloid  variety ;  the  enlargement  of  lymphatic  nodes  was 
trifling.  The  case  to  be  cited  answers  in  most  respects  to 
the  descriptions  of  systemic  lymphomatosis,  but  there 
were  found  two  conditions,  enteritis  and  parasitism, 
which  might  be  responsible  for  sufficient  general  toxemia 
to  stimulate  IjTnph  nodes  and  follicles  to  a  state  of  active 
growth.  Even  accepting  these  two  cases  as  examples  of 
this  group,  it  is  very  plain  that  chronic  systemic  primary 
l>Tnphatic  hyperplasia  is  an  exceedingly  rare  entity  in 
wild  animals.  There  has  been  no  case  resembling 
Hodgkin's  disease  of  man. 

California  Hair  Seal  {Zalophns  calif ornianus) .  Young  6  four 
months  old.  Appeared  to  be  in  good  health,  no  loss  of  flesh  or  activity. 
Diagnosis. — Lymphomatosis.  Fatty  degeneration  of  kidneys.  Em- 
physema. Chronic  follicular  and  catarrhal  enteritis.  Nematodes  in  intes- 
tines. 

External  appearance  good.  Both  lungs  distended  and  are  tense; 
it  seems  almost  like  a  spastic  dilatation — a  simple  emphysema — beneath 
pleural  surface  are  many  minute  petechias.  Lung  mottled  red  and  gray. 
This  color  is  present  on  section.  Lobules  clearly  outlined  and  separable 
with  fingers.  Bronchi  and  vessels  widely  open,  the  latter  containing 
small  amount  of  fluid  blood.  Lymph  nodes — Mediastinal,  tracheal  and 
those  visible  in  neck  are  variously  but  definitely  enlarged,  firm,  irregular 


DISEASES  OF  THE  LYMPHATIC  TISSUES     119 

and  roughly  nodular,  both  in  appearance  and  to  touch.  On  section 
they  are  red  and  gray,  follicular  border  not  preserved  and  merging 
with  medulla;  connective  tissue  visible;  gray  juice  expressed.  Peri- 
cardium normal.  Epicardium,  glistening,  transparent  and  pale.  Heart 
position,  size  and  interior  normal.  Abdomen  contained  no  adhesions  or 
fluid.  Pale  brown  red,  firm  liver  with  smooth  surface  and  sharp  edges; 
markings  not  clear;  section  surface  smooth;  watery  blood  expressed 
from  section.  Gall-bladder  normal,  contents  limpid,  brown.  Common 
bile  duct  patulous.  Somewhat  enlarged  spleen,  soft,  yet  resilient; 
capsule  smooth ;  section  surface  pulp  homogeneous  brown-purple ;  folli- 
cles gray,  slightly  enlarged,  clear,  sharply  outlined;  trabecul^e  faintly 
visible,  more  prominent  where  they  are  near  follicles.  Klidney  normal 
in  size,  shape  and  location,  with  smooth,  pale  brown  surface;  capsule 
smooth,  strips  easily,  section  surface  glistening,  consistency  soft  yet 
resilient ;  thickness  of  cortex  slightly  wide,  of  medulla  normal ;  individual 
lobules  clear  yet  no  increase  of  connective  tissue  between;  pyramids 
quite  pale;  top  of  medulla  a  little  darker;  cortices,  pale,  striae  quite 
faint  and  seem  irregular ;  glomeruli  not  visible.  R.  Adrenal — In  upper 
pole  are  two  cysts  about  1.5  and  .8  em.  across ;  no  scar  at  this  point.  L. 
Adrenal — Wide  pale  purple,  regular  homogeneous  cortex  under  very 
dense  capsule;  medulla  reddish  brown,  homogeneous,  quite  bloody.  In 
left  adrenal  is  a  pale  gray,  fairly  well  outlined  area  between  medulla 
and  cortex  at  upper  pole  about  3  mm.  across. 

Stomach. — Contains  glass  and  stones;  mucosa,  soft,  smooth,  flat, 
yellow  and  pink,  translucent  rugs ;  cardia  about  normal ;  pylorus,  valve 
prominent  but  probably  not  hypertrophied.  Duodenum — Beginning  at 
pylorus  and  extending  through  to  ileum  where  follicles  commence,  intes- 
tine contains  small  amount  white,  creamy  material,  serosa  negative, 
mucosa  smooth  flat,  pale  pink,  translucent.  There  are  small  nematodes, 
probably  uncinaria,  some  of  which  are  attached  quite  firmly.  Here  and 
there  throughout  the  gut  are  bits  of  thickened  mucosa  or  submueosa,  2-5 
mm.  across.  Some  of  these  have  a  tiny  opening  from  which  clear  fluid 
can  be  expressed.  Ileum  shows  smooth  flat  translucent  mucous  mem- 
brane ;  empty ;  individual  follicles  faint  but  discernible.  Colon  reddened 
mucous  membrane,  nematodes  and  little  mucus.  Pancreas,  soft,  normal 
in  size  and  position,  color  pink  gray.  The  lymphatics  of  peritoneal  cavity 
are  all  enlarged;  most  of  retroperitoneal  are  also.  Regional  lymph 
glands  are  also  enlarged.  The  description  given  for  mediastinal  answers 
here.  The  glands  of  mesenteric  stalk  present  chain  of  sausage-like 
masses.  Glands  in  mesentery  are  but  slightly  affected.  Glands  within 
and  without  peritoneum  at  kidney  are  especially  enlarged.  Large  gland 
behind  stomach  and  pancreas  and  in  front  of  upper  end  of  spleen  is 
much  enlarged,  thick  and  roughly  circular;  it  shows  much  fibrosis.  The 
regional  glands  are  distinctly  enlarged  but  proportionately  not  so  much 
as  the  internal  ones.  Smear  of  juice  expressed  from  peritoneal  glands 
shows  large  and  small  lymph  cells  in  about  proper  proportions.  There 
are  a  few  plasma  cells.     No  granular  cells. 


120   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Smear  from  bone  marrow  shows  enormous  number  of  premyeloeytes 
and  mast  cells.  Polynuclears  in  their  early  stages  are  not  numerous. 
Small  gi-oups  of  lymphocytes,  lightly  packed  together,  noted  here  and 
there.    Blood  smears  unsatisfactory. 

JMicroscopical. — Lung  shows  distended  vesicles  with  considerable 
congestion  of  septa  and  in  some  places  red  blood  cells  in  alveoli. 

Kidney. — Glomeruli  contracted,  relatively  anemic ;  capsule  negative ; 
epithelium  throughout  in  state  of  granular  degeneration  probably  fatty ; 
connective  tissue  not  increased. 

Intestines. — Show  slight  hyaline  change  in  muscle  fibres  and  nodular 
degeneration  and  disappearance  of  protoplasm,  leaving  bare  nuclei 
and  outline  of  the  cells;  submucosa  loose;  mucosa  shows  infiltration  of 
round  and  plasma  cells,  diapedesis,  degeneration  and  desquamation  of 
epithelium.  Section  of  ileum  shows  a  hyperplasia  of  lymph 
follicles  with  active  centre  in  which  large  endothelial  cells  pre- 
dominate. Small  swellings  in  submucosa  consist  of  chronic  granulation 
tissue,  but  there  is  no  foreign  body  and  no  evidence  of  preexisteuee 
of  a  lymph  follicle. 

Lymph  nodes  loosely  arranged  but  in  places  solidly  small  lymphoid ; 
chords  and  sinuses  contain  chiefly  small  lymph  cells  in  which  are  some 
mast  and  plasma  cells;  follicles  small  and  loose,  and  centres  contain 
chiefly  large  lymph  and  plasma  cells;  connective  tissue  not  increased 
yet  there  are  some  fibroblasts  in  follicles. 

Spleen. — ShoAvs  almost  entirely  hemoiThagic  pulp ;  blood  destruc- 
tion not  active;  follicles  large  and  made  up  of  about  equal  mixture 
of  small  lymph,  plasma  and  large  lymph  cells ;  connective  tissue  promi- 
nent probably  because  of  excess  of  blood  around  it. 

Local  Hyperplasias. 

In  this  group  and  touching  the  purely  hyperplastic, 
come  the  hypertrophies  of  the  lymphadenoid  tissue  of 
the  nasopharynx ;  this  can  be  disposed  of  by  stating  that 
no  true  chronic  hypertrophies  have  been  seen.  With  most 
cases  of  gastroenteritis  in  ungulates,  and  many  in  car- 
nivores, there  is  a  swelling  and  redness  of  the  tonsillar 
region  but  an  exudative  or  necrotizing  process  does 
not  occur. 

Thymus. 

The  thymus  is  rarely  visible  in  our  specimens  but 
when  found  has  usually  been  normal.  An  increase  in  size 
and  a  decrease  of  consistency  of  this  organ  is  noted  in 
marasmus  (inanition)  from  failure  of  care  of  the  young 


DISEASES  OF  THE  LYMPHATIC  TISSUES     121 

by  the  parent,  but  so  far  as  can  be  determined  this  has 
not  been  adequate  to  cause  tracheal  compression.  In  a 
few  cases  of  rachitis  in  the  canines,  the  organ  is  large  and 
pale  along  with  the  rest  of  the  lymphatic  system.  In  one 
of  the  tumors  of  the  mediastinum  to  be  discussed,  the 
suspicion  arose  that  the  growth  originated  in  the  thjTiius ; 
adenomata  and  sarcomata  have  been  described  in  the 
lower  animals. 

Tuberculosis. 

Tuberculosis  of  the  superficial  lymph  glands  is  rare 
as  an  independent  lesion  in  the  lower  animals.  Dr.  C  Y. 
White  was  fortunate  in  seeing  a  monkey  with  a  chain  of 
fibrocaseous  nodes  in  the  cervical  region,  upon  which 
before  death  he  offered  the  suggestion  that  it  was  of 
tuberculous  nature.  In  Primates  almost  all  drainage 
glands  exhibit  some  miliary  or  caseous  process.  One 
monkey  rejected  upon  the  tuberculin  test  had  what  was 
apparently  a  primary  lesion  in  the  glands  at  the  tracheal 
bifurcation.  In  the  Ungulata,  lymphatic  tuberculosis 
assumes  two  forms,  the  caseous  and  the  cellular.  The 
fonner  is  generally  understood  while  the  latter  is  more 
uncommon.  It  is  occasionally  seen  in  the  ''fungous 
tubercle"  of  cows,  but  we  have  seen  it  in  deer  and  in 
another  order,  Carnivora.  In  the  latter,  tuberculosis  being 
uncommon,  examples  in  the  Ijmiph  nodes  were  noted  but 
twice,  once  caseous  and  once  solid ;  this  latter  was  made 
up  of  firm,  homogeneous  yellow  pink  masses  of  glands  in 
the  mediastinum,  shomng  under  the  microscope  solidly 
packed  epithelioid  and  giant  cells. 

Lymphatic  tuberculosis  in  the  birds  is  rare ;  only  one 
case  is  recalled  (unfortunately  record  cannot  be  found) 
as  small  yellow,  discrete  firm  nodules  in  the  mediastinum 
and  neck.  The  minute  picture  was  of  a  solid  arrangement 
of  large  vacuolated  mono-  and  polynucleated  cells  which 
were  so  packed  with  bacilli  that  the  preparation  could 
not  be  decolorized. 


122   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

TUMOES. 

The  neoplastic  enlargements  are  represented  by  a 
IjTnphosarcoma  of  the  mediastinum  in  a  Dorcas  Goat; 
there  were  secondary  growths  in  the  liver,  the  kidney  and 
several  isolated  lymph  glands.  Secondary  growths  from 
original  tumors  not  in  the  lymphatic  system  are  curiously 
rare,  certainly  much  less  frequent  than  is  found  in  the 
human  being.  In  ninety-two  tumors  which  are  known  to 
give  metastases,  only  three  involved  the  lymphatic  glands 
and  only  one  of  these  could  be  considered  as  involving 
nodes  not  in  the  ordinary  drainage  pathway. 

The  Spleen. 

The  spleen  is  an  organ  whose  exact  position  in  the 
scheme  of  things  remains  mystifying  through  the  whole 
series  of  animals.  Its  functions  have  been  arrived  at 
largely  by  exclusion,  somewhat  by  conjecture  and  specu- 
lation, while  the  acceptably  proven  duties  are  indeed  few. 
Although  this  is  not  the  place  to  enter  into  a  discussion  of 
all  the  points  at  issue  concerning  the  anatomy  and  func- 
tion of  the  organ,  a  few  facts  might  be  mentioned  of 
comparative  and  perhaps  pathological  value.  While  the 
spleen  has  been  looked  upon  always  as  the  origin  of  blood 
cells  in  the  embryo,  cases  in  man  are  on  record  where  no 
spleen  was  discovered,  a  condition  suggesting  that  its 
absence  is  not  incompatible  with  life ;  that  such  is  the  case 
is  obvious  for  experimental  or  therapeutic  removal  of 
the  organ  is  well  known. 

It  was  suggested  by  Virchow  that  at  times  the  spleen 
assumes  the  duty  of  the  bone  marrow  and  the  swelling  of 
the  viscus  in  certain  anemias  seems  to  support  this  state- 
ment. However,  there  is  no  physical  relationship  between 
the  size  of  the  animal  and  that  of  the  spleen,  nor  between 
the  available  quantity  of  bone  marrow,  the  obvious  rich- 
ness of  circulating  blood  and  amount  of  lymphatic  tissue. 
This  is  well  shown  in  the  water  mammals  which  have  a 
great  deal  of  blood  in  vessels  and  viscera  yet  their  spleen 


DISEASES  OF  THE  LYMPHATIC  TISSUES     123 

is  relatively  smaller  than  birds  of  somewhat  comparable 
size  (the  ostrich,  for  example).  The  Carnivora  and 
Eodentia  among  the  mammals,  have  quite  large  spleens 
proportionately,  while  comparable  birds,  Accipitres  and 
Striges,  have  relatively  small  ones,  and  an  absolutely 
small  amount  of  marrow. 

Another  anomalous  example  of  the  function  of  this 
tissue  is  found  in  the  hematopoiesis  of  birds.  While  there 
seems  to  be  some  evidence  that  in  the  spleen  and  liver 
red  blood  cells  may  be  formed,  there  is  perhaps  at  most 
times  sufficient  bone  marrow  to  keep  the  blood  cells  at  a 
proper  number.  There  is  no  inverse  relation  between  the 
amount  of  bone  marrow  and  that  of  splenic  bulk ;  that  is 
in  diving  birds  with  their  hollow  bones  the  latter  is  no 
greater  than  in  gallinaceous  birds  which  have  abundant 
marrow  in  all  the  skeleton. 

It  has  been  suggested  that  the  spleen  supplies  an 
activator  for  pancreatic  ferments.  This  receives  a  sort  of 
support  from  the  greater  size  of  the  organ  in  carnivores 
and  smaller  relative  size  in  ungulates,  since  in  the  former 
concentrations  of  digestive  enzymes  are  more  often 
needed.  Because  of  the  greater  excretion  of  iron  in 
splenectomized  animals  or  those  being  starved,  Fischer 
has  thought  that  the  spleen  has  some  power  to  metabolize 
this  element ;  this  receives  some  support  from  the  state- 
ments concerning  the  size  of  this  organ  in  the  carnivorous 
orders.  From  experimental  studies  it  appears  that  some 
relation  exists  between  the  lymphocytes  and  resistance  to 
implanted  tumors.  If  one  apply  this  idea  to  the  amount 
of  lymphatic  tissue  and  the  incidence  of  spontaneous  new 
growths,  it  is  found  that  no  definite  relationship  is  discov- 
erable either  in  terms  of  size  of  spleen  or  richness  in 
lymphatic  nodes.  Carnivores,  rodents  and  marsupials 
show  a  high  tumor  incidence  and  have  a  good  lymphatic 
supply.  Aves,  on  the  other  hand,  with  a  poor  nodal  appa- 
ratus and  a  variable   splenic  bulk,   show  many  fewer 


124   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

tumors  than  do  the  mammals.  The  mononuclears  of  avian 
blood  rise  normally  as  high  as  60  per  cent.,  of  which 
40-45  per  cent,  are  of  the  small  size. 

The  size  of  the  spleen  is  subject  to  great  variation 
not  only  within  orders  but  actually  within  genera ;  it  even 
seems  that  one  sometimes  sees  two  or  three  members  of 
a  species  kept  in  the  same  enclosure,  maybe  with  the  same 
disease,  maybe  without  any  obvious  disease,  yet  with 
definite  variation  in  the  size  of  the  organ.  This  irregu- 
larity is  recognized  by  veterinarians  (Hutyra  and  Marek) 
who  ascribe  it  to  some  unknown  disease,  past  or  present, 
and  to  normal  variation.  The  lack  of  uniformity  is  seen 
more  clearly  in  birds  than  mammals.  In  the  former  it 
might  be  due  at  times  to  low  grade  or  inactive  parasitism 
or,  conjecturally,  to  toxins  from  incorrect  diet ;  we  shall 
see  later  that  infection  and  intoxication  seem  to  have  a 
different  effect.  Some  writers  have  mentioned  the  possi- 
bility of  an  idiopathic  splenomegaly  (?),  a  condition 
associated  in  youthful  human  beings,  with  anemia  and 
lymphadenopathy,  and  occasionally  going  over  into  a 
sort  of  leucemia.  The  existence  of  such  a  condition  is 
difficult  to  admit  or  deny;  we  have  met  nothing  which 
could  not  be  aligned  in  some  fairly  well  defined  group. 
Birds  have  a  relatively  larger  spleen  than  mammals  and 
in  addition  the  organ  seems  to  respond  more  actively 
in  infections  or  intoxications,  since  it  may  reach,  under 
active  stimulation,  a  size  ten  times  that  of  the  nor- 
mal organ. 

Enlargements  of  Spleen. 

Acute  enlargements  of  the  spleen,  be  they  of  con- 
gestive or  ''inflammatory"  nature  are  quite  common 
among  the  lower  animals,  a  fact  that  is  recognized  in 
veterinary  medicine.  Congestions  of  the  spleen  are  most 
often  seen  during  acute  infections  and  diseases  of  the 
heart  and  liver.     The  diagnosis  has  been  made  among 


DISEASES  OF  THE  LYMPHATIC  TISSUES     125 

mammals  proportionately  more  than  in  birds,  27  or  1.5 
per  cent,  to  35  to  1  per  cent. 

Eeason  foe  Congestive  Enlakgements. 

Perhaps  this  is  partly  due  to  the  circulatory  anatomy 
since  in  the  former  the  arterial  and  venous  supply  is 
usually  by  one  large  vessel  of  each  kind,  whereas  in  birds 
the  splenic  branch  of  the  coeliac  axis  breaks  up  into  several 
small  arteries  and  the  venous  return  is  accomplished  by 
numerous  venules  some  of  which  reenter  the  posterior 
cava  almost  directly,  others  joining  with  the  mesenteric 
to  form  the  portal;  by  this  arrangement  a  more  elastic 
system  is  assured.  The  same  condition  is  found  when 
analyzing  the  records  of  chronic  passive  congestion;  in 
eighteen  recorded  cases,  fourteen  were  mammals  and  four 
birds.  In  these  cases  the  principal  associated  pathologi- 
cal lesions  were  pleural  and  pulmonary  in  seven,  cardiac 
in  three,  renal  in  three  and  hepatic  in  four.  While  the 
anatomy  of  the  splenic  blood  supply  may  help  to 
explain  the  small  number  of  congestions  in  Aves,  it  will 
not  answer  for  the  inflammations  which  occur  in  large 
numbers  in  this  class;  this  will  be  discussed  in  the 
following  pages. 

Hemorrhage  and  infarction  of  the  spleen  are  not 
very  common,  there  being  eleven  of  the  former  and  ten  of 
the  latter  and  all  occurring  with  the  same  indication  or 
history  of  infectious  disease;  in  one  case,  an  opossum 
{Didelphys  virginiana)  an  injury  probably  caused  a  mas- 
sive hemorrhage  shaped  like  an  infarct.  It  is,  however, 
curious  that  of  the  twenty-one  cases  only  five  occurred  in 
birds,, of  which  only  one  hemorrhage  was  in  the  shape  to 
which  the  term  infarction  is  best  applied.  There  have 
been,  as  one  might  expect,  a  few  cases  of  infected  infarc- 
tion, with  abscess.  It  is  perhaps  worth  noting  that  no 
case  of  hemorrhagic  cyst  or  inspissated  coagulum  has 
been  seen.  Only  one  case  of  rupture  is  recorded  second- 
ary to  acute  splenitis  during  an  acute  septicemia. 


126   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 


Inflammations. 

It  has  been  the  practice  at  this  laboratory  to  classify 
splenitis  in  three  ways,  (1)  acute  diffuse  splenitis — 
general  congestion  with  perhaps  small  hemorrhages, 
swelling  but  retention  of  general  architecture  and  mth  no 
undue  prominence  of  the  follicles;  (2)  follicular  hyper- 
plasia— where  this  is  the  prominent  gross  and  minute 
finding,  the  pulp  being  less  pronouncedly  involved; 
(3)  acute  splenic  tumor — where  the  organ  is  greatly 
enlarged  but  with  loss  of  the  usual  markings.  Perhaps 
this  separation  is  not  warranted  upon  a  strict  etiological 
basis,  and  yet  as  we  shall  see  it  holds  true  fairly  well  in 
the  toxic  and  infectious  lesions..  Moreover,  from  the  fol- 
lowing list  it  is  evident  how  the  various  orders  call  upon 
the  splenic  tissue  in  disease. 

Table  7. 
This  Shows  the  Percentage  of  Various  Forms  of  Splenitis  in  the  Animals  Coming 
to  Autopsy. 


Acute  Diffuse 
Splenitis. 
Per  cent. 

Follicular 
Splenitis. 
Per  cent. 

Acute  Splenic 
Tumor. 
Per  cent. 

7. 

2.5 

2.4 

2.6 

9.2 

SS. 

S3. 

12.5 

3. 
6.5 
5.2 
1. 
2.6 
10. 
7.5 

4. 

6.8 

3. 

.7 

3. 
4.9 
.3 
4.7 
3. 

3. 
2.3 
7.2 
6. 
1.5 
10. 
3.7 

3.4 
6. 
10. 

.7 

1.3 

2. 

1.2 

1. 

Marsupialia 

Pinnipedia     

3.7 

Proboscidea.              

Edentata                          

6.2 

Passeres                            

3.8 

Psittaci 

4.1 

5.2 

2. 

Galli     

4.7 

TT 

Accipitres 

1.6 

GaviaB                    

5. 

Picarise                       

1.3 

Striges                                    

FulicarisG 

Stecanopodes 

For  the  meaning  of  italics  see  foot  note  Table  1. 


'   The  points  to  draw  from  this  chart   are  first  the 
greater  variety  of  lesions  seen  in  the  birds,  which  appar- 


DISEASES  OF  THE  LYMPHATIC  TISSUES     127 

ently  make  greater  use  of  the  organ  in  defence,  and  the 
preponderance  of  acute  diffuse  hyperplasias  in  mammals. 
It  is  noteworthy  that  the  Primates  and  Marsupialia  show 
more  of  the  acute  hypertrophy  of  the  spleen  going  under 
the  name  of  tumor.  It  would  be  interesting  and  valuable 
to  be  able  to  discover  exactly  what  determines  and  con- 
stitutes the  conditions  usually  termed  hyperplasia  and 
inflammation  of  the  spleen  and  an  attempt  was  made  in 
this  direction  by  tabulating  the  data  from  Table  7  in 
terms  of  each  diagnosis,  classifying  these  latter  also  as 
infectious  and  toxic.  The  results  of  this  effort  are  not 
conclusive,  and  while  they  permit  of  some  discussion  of 
the  lesions,  do  not  allow  finished  conclusions.  The  figures 
obtained  by  study  are  not  illuminating.  One  can  state, 
however,  that  in  infections,  either  specific  or  not  specific, 
more  elements  of  the  spleen  were  engaged  in  the  process 
than  when  the  condition  did  not  resemble  a  communicable 
disease,  but  might  be  called  toxic.  Under  the  former 
condition  the  diagnosis  of  splenitis  or  acute  tumor  pre- 
dominated, while  under  the  latter  follicular  hyperplasia 
is  more  often  recorded.  Anatomical  alterations  in  these 
two  groups  are  described  in  the  definition  given  in  a 
previous  paragraph  and  deserve  no  special  discussion 
except  in  so  far  as  they  concern  the  changes  in  the  avian 
splenic  tumor  when  under  magnification. 

Microscopically  there  is  a  total  loss  of  the  relations  in 
the  acute  splenic  tumor  of  birds,  the  chords  and  follicles 
being  replaced  by  a  rather  regularly  arranged  mass  of 
small  and  large  mononuclears,  granular  cells  of  the  circu- 
lating types  and  red  blood  cells ;  pigment  always  seems 
increased.  Endothelial  cells  do  not  take  part  in  the 
general  mass  of  the  organ  but  along  lymph  and  blood 
vessels  their  bulk  and  number  are  increased  so  that  if  the 
packing  be  not  too  dense  one  may  find  double  lines  of  these 
cells  passing  through  the  hyperplastic  tissue.  :There  is, 
however,  a  group  of  seven  cases  (six  birds)  to  which  the 
term   endothelial   hyperplasia   in   the    spleen  has   been 


128   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

applied.  All  of  them  show  an  unusual  prominence  of  the 
vascular  linings  and  of  the  follicular  centres  and  peri- 
vascular areas ;  besides  this  there  is  a  moderate  general 
hypei*plasia.  An  examination  of  the  historj^  and  autopsy 
results  in  the  birds  indicated  that  they  had  all  suf- 
fered with  some  rather  protracted  intoxication  and 
showed  a  moderate  anemia  (four  of  the  spleens  were 
well  pigmented). 

The  more  chronic  changes  of  this  organ,  be  they 
moderate  or  of  a  grade  to  which  one  must  apply  the  term 
fibrosis,  are  rather  uncommon  when  one  considers  the 
number  of  animals  with  prolonged  infection,  anemia, 
skeletal  diseases  and  hepatic  cirfhoses ;  these  are  the  con- 
ditions that  take  a  prominent  place  in  the  associated 
pathology  and  history.  There  is  no  essential  difference  in 
the  organ  throughout  the  animal  kingdom,  and  one  may 
find  as  far  down  as  the  struthious  birds  analogies  to  the 
processes  of  domestic  and  human  animals. 


Special.  Subjects — Amyloid. 

Amyloid  changes  in  the  spleen  have  occurred  in 
the  follomng  orders:  Primates,  Carnivora,  Rodentia, 
Marsupialia,  Passeres,  Galli,  Gaviae,  and  Anseres, 
fourteen  cases  in  all.  This  infiltration  is  usually 
ascribed  to  long  continued  suppuration  or  chronic 
infection,!  every  one  admitting,  however,  that  once  in  a 
great  while  a  case  is  seen  wherein  no  cause  can  be  dis- 
covered. In  this  laboratory  we  have  a  high  percentage 
of  cases  without  adequate  accompanying  pathology 
so  that  we  have  called  six  of  the  above  cases 
primary  or  idiopathic.  The  gross  appearance  of 
the  spleen  is  in  all  these  cases  that  of  an  enlarged, 
firm,  homogeneous  body  without  the  sago  spots  usually 
described  for  this  change.  \  Microscopically  the  infiltra- 
tion occurs  first  in  the  vessel  walls,  thence  spreading  to 
the  sinus  walls  and  reticulum.    Joest  in  discussing  this 


DISEASES  OF  THE  LYMPHATIC  TISSUES     129 

infiltration  (1),  states  that  it  is  comparatively  rare  among 
the  domestic  animals,  where  it  may  appear  mider  the 
usual  conditions  or  as  a  primary  affection;  it  seems  at 
times  to  follow  unsuitable  feeding.  According  to  this 
author,  the  ''sago"  type  is  more  common  than  the 
''bacony;"  this  has  not  been  our  experience.  The  most 
conspicuous  example  is  given  here. 

European  Badger  {Meles  meles)  Adult    9.  Gradual  failure  for  two 
weeks. 

Diagnosis. — Amyloidosis.      Chronic      parenchymatous      nephritis. 
Emphysema  of  lungs. 

Muscles  atrophic;  fat  scanty.  Heart  is  dilated  and  muscle  is 
red  brown.  Aorta  is  jaundiced  and  there  is  a  small  patch  of  atheroma 
near  the  anterior  leaflet  of  aortic  valve  and  about  5  mm.  above  it. 
Abdomen  contains  a  slight  amount  of  clear  fluid,  no  adhesions.  Liver 
is  normal  in  size,  smooth  surface,  sharp  edges,  firm  consistency,  yellow 
color.  The  section  surface  is  glistening,  smooth  and  moist.  The  spleen 
is  very  hard,  greatly  enlarged  (25  x  6  x  2  cm.)  and  lies  across  the  abdo- 
men over  the  intestines.  It  is  pale  red  and  very  bacony.  In  iodin  the 
tissue  stains  a  mahogany  brown.  The  kidney  is  normal  in  size.  (5^2  x  3 
cm.).  The  capsule  is  smooth  and  strips  easily  leaving  a  smooth,  yellow 
surface.  Organ  is  firm.  The  section  surface  is  glistening,  has  a  relatively 
narrowed  cortex  and  relatively  wide  medulla.  The  glomeruli  are  barely 
visible  but  stain  a  mahogany  brown  in  iodin.  The  adrenals  are  12  xlO  x  4 
mm.  The  cortex  is  wide,  dull  yellow  and  regular.  The  zone  beneath 
fades  into  the  cortex  although  rather  abruptly.  The  centre  of  both 
glands  is  occupied  by  an  irregular  pale  yellow,  sharply  outlined  nodule 
suggestive  of  solid  medulla,  tubercle  or  tumor.  The  zone  between  cortex 
and  this  is  gray,  irregular  and  firm.  This  is  practically  a  normal  figura- 
tion. The  stomach  is  empty,  mucosa  apparently  normal.  Intestines 
were  not  opened  but  serosa  seems  normal ;  when  opened  after  Kaiserling 
fixation  they  seem  normal.  The  pancreas  is  normal  in  size,  firm  and 
pale.  Lymphatics  of  the  mesentery  are  slightly  enlarged,  soft,  homo- 
geneous pale  yellow. 

Histological  Notes. — Lung  seems  somewhat  atrophic  and  there  are 
some  vesicular  ruptures.  One  small  patch  of  amyloid  found  in  blood 
vessel  wall.  Liver  shows  slight  capillary  congestion  and  granularity 
of  parenchyma.  There  is  a  marked  blood  vessel  amyloid  deposit  not 
only  in  interlobular  spaces  but  in  intralobular  capillaries.  Spleen,  no 
trace  of  splenic  tissue  recognizable  in  section.  It  is  composed  of  more 
or  less  eosin-staining  material  surrounding  single,  or  small  collections 
of  round  or  plasma  cells.  Kidney  shows  marked  amyloidosis  of  glomeruli 
and  slight  deposit  in  blood  vessel  walls  and  in  the  increased  connective 
tissue.  There  is  a  general  moderate  fibrosis;  irregular  tubules;  low 
(1)    Lubarsch-Ostertag,  Ergeb.  aus  der  Allg.  Path.,  1908. 


130   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

epithelium  and  hyaline  casts.  Heart  muscle  fibres  are  small  and  stain 
deeply.  No  amyloid  in  section.  No  pigmentation  although  section  is 
suggestive  of  brown  atrophy.  Adrenal  is  practically  negative.  There 
seems  to  be  slightly  more  connective  tissue  than  noimal  but  parenchyma 
may  be  considered  normal.  At  one  place  in  the  cortex  there  are  some 
structures  of  deep  layer  included  in  vesicular  layer.  This  seems  like 
a  structural  malformation. 

Necroses. 

Focal  necroses  of  the  spleen  affecting  chiefly  follicular 
centres  but  also  chords,  are  not  at  all  uncommon  in  avian 
spleens,  especially  where  parasitism  occurs,  not  only  with 
hemic  protozoa  and  embryos,  but  also  ^\ith  intestinal  or 
visceral  nematodes  and  trematodes. 

Spleen  in  Anemia. 

The  spleen  in  the  anemias  shows  much  less  definite 
change  than  one  would  expect.  In  the  secondar^^  form 
of  anemia  among  mammals  one  finds  a  slight  excess  of 
pigmentation  and  an  occasional  fibrosis  but  often  the  size 
of  the  organ  is  recorded  as  normal.  When  the  impover- 
ished blood  seems  secondary  to  skeletal  degenerations 
there  is  a  diffuse  or  follicular  enlargement.  In  the  birds, 
on  the  other  hand,  there  is  nearly  always  some  grade  of 
enlargement  which  is  due  in  the  well  studied  examples  to 
a  richness  of  blood  cells.  There  may  be  a  slight  increase 
in  follicles,  but  these  bodies  are  usually  small  and  solid. 
In  two  instances  a  prominence  of  large  endothelial  cells 
was  discovered.  Pigment  is  seldom  increased,  but  it 
may  be  very  excessive. 

In  the  primar}^  anemias  little  more  than  the  above  is 
to  be  found.  Fibrosis  is  more  evident  and  perhaps  pig- 
mentation less  so,  but  the  variations  are  more  of  degree 
than  kind.  The  spleen  in  leucemia  can  only  be  discussed 
upon  the  case  already  reported  {q.  v.). 

Spleen  in  Hepatic  Fibroses. 

The  fibroses  of  the  liver  are  not  infrequently  associ- 
ated with  some  enlargement  of  the  organ  under  discus- 
sion.   At  this  laboratory  hepatic  cirrhoses  are  divided 


Fig.  7. — EUROPEAN  BADCl.R  IM1.I.1.>  Ml.l.L.i).  PRIMARY  AMYLOIDOSIS.  THE  ENOR- 
MOUSLY ENLARGED  SPLEEN  IS  SEEN  LYING  ACROSS  THE  STOMACH  BELOW  THE  LIVER, 
WHICH   ORGAN  IS  EXCEEDED  IN  SIZE  BY  IT. 


DISEASES  OF  THE  LYMPHATIC  TISSUES     131 

into  Portal,  Biliary,  Fatty,  Perilobular  and  Vascular. 
By  a  study  of  the  spleens  in  these  cases  a  few  facts  have 
been  obtained.  In  mammalian  portal  cirrhoses  there  is 
usually  a  very  moderate  but  definite  increase  in  the  size 
of  the  spleen  due  to  connective  tissue  increase  in  the 
trabeculae  and  reticulum,  with  very  small  compact  fol- 
licles. Among  the  birds  the  organ  seldom  shows  more 
that  a  moderate  congestion  and  diffuse  hyperplasia.  In 
the  mammalian  biliary  cirrhoses  there  is  almost  without 
exception  a  definite  enlargement  due  to  fibrosis  and  fol- 
licular hyperplasia.  Among  the  birds  the  process  is  very 
far  from  uniform,  there  being  just  as  often  no  change,  as 
a  congestion  and  hyperplasia,  or  as  a  mild  fibrosis ;  it  is 
notable,  however,  that  pigmentation  is  commonly  met  in 
this  class.  No  noteworthy  change  is  met  \vith  in  the 
spleen  of  fatty  hepatic  cirrhoses.  We  have  no  record 
of  cyanotic  induration  of  the  spleen  accompanying  a 
similar  condition  in  the  liver.  When  the  condition  of 
perilobular  hepatic  fibrosis  has  been  met  a  distinct 
increase  of  the  splenic  pigment  is  usually  found.  In 
reviewing  the  facts  in  this  paragraph  one  is  forced  to  the 
conclusion  that,  with  the  exception  of  the  frank  infective 
cases,  there  is  no  definite  relation  of  the  splenic  changes 
to  those  in  hepatic  cirrhosis,  a  fact  made  perhaps  the 
more  significant  in  view  of  the  idea  held  in  some  quarters 
that  the  primary  change  in  this  pathological  process 
occurs  in  the  spleen. 

Perisplenitis. 

Perisplenitis  of  an  acute  or  fibrosing  variety  as  a  part 
of  peritonitis  or  as  the  result  of  an  injury  to  the  splenic 
region,  has  been  encountered  on  numerous  occasions  and 
offers  nothing  worthy  of  comment,  but  the  so-called 
''sugar-icing"  spleen,  associated  with  perihepatitis  and 
general  peritoneal  thickening,  and  with  an  obscure  rela- 
tion to  tuberculosis  has  not  been  seen.  As  we  shall  learn 
later  peritoneal  or  massive  intestinal  tuberculosis  is  not 


132   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

common  in  the  lower  animals.  A  diffuse  fibrous  thick- 
ening of  a  pearl  gray  glistening  appearance,  has  occur- 
red in  five  monkeys  harboring  the  Filaria  gracilis  in 
the  peritoneum. 

Tuberculosis. 

In  tuberculosis  of  this  organ  the  capsule  is  almost 
invariably  thickened  over  the  nodules,  but  not  over  the 
whole  organ  and  rarely  in  the  diffuse  variety.  There  is 
almost  always,  however,  in  this  prolonged  infectious  dis- 
ease some  reaction  on  the  part  of  the  spleen,  particularly 
when  several  other  viscera  are  involved.  In  such  cases 
a  low  grade  of  general  hyperplasia  and  fibrous  tissue 
increase  is  found.  This  is  especially  true  in  the  Primates 
and  Carnivora  in  long  continued  pulmonary  tuberculosis, 
and  it  is  in  this  form  that  one  does  not  often  see  tubercu- 
lous lesions  proper  in  the  spleen  itself.  Tuberculosis  of 
the  spleen  is  not  very  common  in  adult  human  beings  and 
domesticated  animals,  while  in  the  young  it  is  seen  in  a 
fairly  high  percentage  of  cases.  In  the  wild  animals  this 
organ  seems  much  more  susceptible  to  the  settling  of 
tubercle  bacilli  as  will  be  attested  by  the  facts  to  be  cited, 
and  yet  there  are  some  conspicuous  exceptions  to  this 
statement.  Just  why  certain  groups,  or  orders  should 
show  splenic  tuberculosis  very  frequently  while  others 
fail  to  do  so  is  far  from  clear. 

The  appearance  of  the  lesion  is  fairly  comparable 
throughout  the  animal  kingdom,  variations  among  the 
Aves  being  chiefly  in  the  more  frequent  occurrence  of 
the  diffuse  tuberculous  splenitis.  Miliary,  caseous  and 
conglomerate  masses  occur  separately  or  together  and 
without  very  distinct  relation  to  other  pathologi- 
cal involvements. 

The  relative  incidence  of  tuberculosis  in  the  spleen  is 
set  forth  in  the  section  devoted  to  this  infection,  but  may 
be  mentioned  here  in  a  general  way.  The  organ  is  about 
equally  susceptible  to  the  disease  in  mammals  and  birds 


DISEASES  OF  THE  LYMPHATIC  TISSUES     133 

according  to  the  figures,  but  the  high  percentage  for  the 
former  is  due  to  the  frequency  with  which  tuberculosis  is 
met  in  the  Primates.  With  this  order  deducted  there  is 
no  doubt  that  the  avian  spleen  has  a  greater  vulnerability 
for  the  tubercle  bacillus  than  has  the  mammalian  organ. 
Rodentia  as  a  mammalian  order  stand  next  to  the 
Primates,  whereas  the  Camivora  and  Ungulata  relatively 
seldom  show  splenic  tuberculosis.  Among  the  Aves  one 
finds  that  Columbae,  Galli  and  Accipitres  have  the  highest 
splenic  susceptibility,  but  beyond  this  one  hardly  dare 
venture  because  of  the  irregularity  in  the  number  of 
specimens  seen  at  autopsy.  Suffice  it  to  say  that  a  greater 
number  of  cases  of  tuberculosis  are  seen  in  the  spleen 
of  the  class  for  which  the  intestinal  origin  of  tuberculosis 
appears  most  important. 

In  the  cases  of  actinomycosis  and  its  congeners,  such  as 
the  Kangaroo  maxillarj^  mycosis,  which  we  have  seen  at 
the  Garden,  no  specific  changes  have  been  seen  in  the 
spleen,  unless  a  single  or  double  nodule  of  the  same  variety 
as  the  original  focus  may  be  considered  specific.  Such  was 
found  in  two  cases,  one  a  tapir  Avith  low  grade  inactive 
actinomycosis,  the  other  a  kangaroo  with  maxillary  strep- 
tothricosis.  Two  deer  came  to  autopsy  with  a  gross 
picture  suggestive  of  anthrax;  no  bacilli  could  be  found 
in  the  semifluid  spleen,  but  a  member  of  the  hemorrhagic 
septicemia  group  w^as  found.  The  latter  group  of  infec- 
tions has  been  fairly  well  represented,  although  not  in 
epizootic  form,  and  the  almost  invariable  splenic  change 
has  been  that  already  discussed  as  diffuse  splenitis. 
Three  secondary  tumors  are  recorded,  two  cancers  and 
one  renal  adenoma. 


SECTION  VI 

THE  RESPIRATORY  SYSTEM  AND  ITS 
RELATED  STRUCTURES 

It  is  customary  to  divide  the  descriptions  of  normal 
and  diseased  conditions  of  the  upper  entrance  to  the  body 
into  respiratory  and  alimentary  parts,  the  nose,  naso- 
pharynx and  larynx  belonging  to  the  former,  the  mouth, 
buccal  cavity  and  phar^mx  to  the  latter.  As  a  matter  of 
fact  they  can  for  most  purposes  be  considered  as  the 
structures  contained  in  the  anterior  head  and  furthermore 
their  pathological  states  are  more  often  followed  by  exten- 
sions into  or  implications  of  the  respiratory  organs 
proper  than  of  the  alimentary  tract.  From  a  comparative 
standpoint  the  incidence  of  specific  infectious  diseases  and 
of  the  involvement  of  accessory  nasal  sinuses  present  the 
most  interesting  subjects.  There  are  several  infections, 
believed  to  be  specific,  observed  among  domestic  mammals 
and  birds  but  their  actual  indi\dduality  has  hardly  been 
unexceptionally  proved.  This  refers  to  the  communicable 
rhinitis  of  cows,  pigs,  rabbits,  and  birds,  especially 
parrots,  the  follicular  catarrh  of  horses,  and  croupous 
nasopharyngitis,  all  of  which  have  been  ascribed  to  a  par- 
ticular virus,  without  finished  evidence  in  many  instances. 
Some  of  these  diagnoses  doubtless  cover  or  are  confused 
wdth  the  early  sj^mptoms  and  signs  of  the  disease  of  pro- 
tean manifestations,  distemper,  and  indeed  the  Bact. 
septicus  and  relatives  of  the  bird  cholera  organisms  are 
reported  as  being  responsible  for  them.  No  intention  of 
excluding  well  recognized  entities  like  bird  diphtheria, 
foot  and  mouth  disease  or  influenza,  exists.  I  shall  refer 
below  to  small  groups  of  epizootics  which  do  not  conform 
strictly  with  word  pictures  drawn  by  Hutyra  and  Marek, 
Moore,  or  Ward  and  Gallagher. 

134 


THE  RESPIRATORY  SYSTEM  135 

Rhinitis,  Sinusitis. 

The  nature  of  inflammations  of  the  nasopharynx  sug- 
gests at  once  that  there  may  be  some  anatomical  reason 
for  their  distribution  and  character.  A  general  review  of 
the  anatomy  of  the  mammalian  and  avian  nasopharynx 
reveals  the  relatively  greater  space  in  the  former,  espe- 
cially in  the  passage  from  the  nose  to  the  pharynx,  and 
emphasizes  the  exposure  of  the  opening  of  the  upper 
larynx  in  the  bird,  lying  as  it  does  in  the  posterior  part  of 
the  tongue  and  surrounded  by  the  constrictor es  glottidis. 
A  dissection  of  the  accessory  nasal  sinuses  exposes  the 
relatively  large  size  of  these  spaces  in  the  lower  mammals, 
and  the  capacious  openings  into  the  nasal  cavities.  (1)  In 
the  Primates  and  Lemures  the  anatomy  more  closely 
resembles  that  of  man,  the  sinuses  being  relatively  smaller 
and  the  communicating  passages  narrower.  In  the  bird 
on  the  other  hand,  while  the  sinuses  may  be  extensive  in 
some  they  are  usually  small,  yet  in  all  the  communication 
with  the  turbinate  area  is  by  a  narrower  slit  or  tortuous 
canal,  frequently,  as  in  Galli,  running  from  below  upward 
into  the  maxillary  sinuses.  The  extent  of  the  turbinate 
and  the  richness  in  mucosa  is  probably  greater  in  all 
mammals  than  in  birds ;  certainly  this  seems  true  of 
Carnivora  and  Ungulata  versus  Accipitres  and  Galli. 

If  the  seriousness  of  a  rhinitis  be  dependent  upon  the 
extent  of  involvement  of  the  sinuses  and  the  blocking  up 
of  their  outlets  it  would  be  expected  that  the  variety  of 
animal  having  the  smallest  drainage  channels  would  show 
the  greatest  evidence  of  these  diseases.  Our  records 
would  indicate  that  32  birds  (.96  per  cent,  of  the  autopsies 
upon  Aves)  had  rhinitis  whereas  only  7  mammals  (.39 
per  cent,  of  autopsies  on  this  class)  presented  the  con- 
dition. Extension  to  the  sinuses  occurred  in  only  one- 
third  of  each  of  these  figures,  a  complication  which 
in  turn  produced  generalized  infection  more   often  in 

(1)  Vide  Sisson's  Veterinary  Anatomy  and  Owen's  Anntomy  of 
the  Vertebrates. 


136   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

maminals  than  in  birds  as  4  is  to  3.  These  figures  are 
perhaps  too  small  for  conclusions  but  it  would  seem  that 
rhinitis  occurs  more  often  in  birds  with  their  small  sinuses 
and  channels  wliile  sinusitis  and  general  infection  occur 
more  often  in  mammals  with  their  large  sinuses  and  exten- 
sive turbinate  apparatus.  The  most  conspicuous  orders 
represented  are,  in  line  of  numbers  Anseres  12,  Psittaci 
7,  Accipitres  5,  Carnivora  3.  Nine  of  the  twelve  water- 
fowl were  part  of  an  epizootic  which  mil  be  discussed 
under  specific  diseases. 

Baeteriologically  the  mammalian  cases  that  have  been 
worked  out  were  due  to  Streptococcus  pyogenes  in  several 
instances,  including  the  generalized  cases,  and  to  a 
mixture  of  streptococci,  golden  staphylococci  and  mem- 
bers of  the  colon-aerogenes  group.  In  one  case  in  a 
tapir  a  member  of  the  B.  septicus  group  was  found. 
Moulds  were  discovered  in  three  avian  cases  and  filaria  in 
one.  No  pentastomum  or  oestrus  has  been  discovered.  In 
thirteen  instances  the  lungs  have  been  involved,  appar- 
ently secondary  to  the  nasopharyngeal  disease. 

There  have  been  two  small  outbreaks  of  an  acute  non- 
specific infection — that  is  not  suggestive  of  cholera,  psit- 
tacosis or  infectious  enteritis — among  the  parrots  in 
which  during  a  short  time  4  and  6  parrots  died  mth  naso- 
pharyngosinusitis  as  the  prominent  lesion.  One  outbreak 
was  studied  baeteriologically  without  definite  result. 
There  was  no  uniform  internal  pathology  unless,  in  one 
outbreak,  congestion  of  the  cerebellum  may  be  mentioned. 
Fowl  diphtheria  and  its  associated  condition  from  which 
a  satisfactory  separation  has  not  been  accomplished, 
epithelioma  contagiosum,  has  happily  given  us  little  con- 
cern, so  that  it  is  not  possible  to  record  any  instructive 
facts  upon  its  cause  or  differential  diagnosis.  There  was 
recorded  in  the  1911  Report  of  the  Society  the  occurrence 
of  two  fatal  cases  in  cassowaries  from  which  it  was  pos- 
sible to  isolate  the'B.  columbarum  and  one  bird  with  the 
same  clinical  appearances  whose  recovery  seemed  to  be 


THE  RESPIRATORY  SYSTEM  137 

due  to  the  use  of  human  diphtheria  antitoxin.  In  light  of 
more  inf  onnation  and  consideration  of  the  accepted  varia- 
bility of  this  disease,  it  is  possible  that  tliis  bird  may  have 
recovered  without  the  injections  or  with  the  use  of  normal 
serum.  Nowadays  it  is  possible  to  obtain  antiroup  serum 
which  is  stated  by  Blair  of  New  York  to  be  efficacious. 
The  disease  has  been  observed  in  a  wild  turkey  and  an 
Abyssinian  Ground  Hornbill,  beside  the  two  struthious 
birds  mentioned  above. 

Mycosis. 

Mycotic  disease  of  the  nasopharynx  seldom  restricts 
itself  to  this  cavity,  usually  extending  by  continuity  to  the 
esophagus,  or  by  inspiration  to  the  lungs  whence  it 
spreads  to  the  air  sacs.  This  condition  of  the  upper 
passages  has  occurred  here  only  in  Psittaci  and  Accipitres 
although  it  is  reported  by  veterinarians  as  occurring  in 
Anseres  and  Struthiones.  In  the  first  order  four  birds 
were  affected,  two  showing  extension  to  the  esophagus 
and  a  like  number  having  pulmonary  and  serous  mem- 
brane involvement.  These  cases  were  all  due  to  asper- 
gillus  whereas  those  next  to  be  mentioned  were  caused 
by  an  oidium  close  to  the  ' '  albicans  ' '  variety.  In  four 
Mississippi  kites  the  prominent  changes  were  found  in  the 
pharynx  and  esophagus  down  as  far  as  the  proventricle 
with  only  a  few  rather  trifling  lesions  in  the  nasal  area. 
Infiltrative  and  necrotizing  processes  characterized  the 
action  of  the  oidium  wliile  that  exerted  by  the  aspergillus 
was  more  superficial  and  extensive.  In  one  case  of  a 
parrot  the  whole  nasal  ca\ity  was  completely  filled  with  a 
yellow  gray  exudate  whereas  the  esophageal  wall  of  the 
kites  was  thoroughly  infiltrated  by  a  gray  brown,  friable, 
necrotic  mass.  Attempts  at  treatment  were  made  in  the 
case  of  the  latter,  using  potassium  chlorate  and  saline 
solution  on  cotton  swabs.  The  result  was  entirely  nega- 
tive and  the  applications  seemed  to  have  no  effect  upon  the 
course  of  the  infection. 

10 


138   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

There  are  on  our  records  in  addition  to  the  above, 
several  cases  of  necrotizing  processes  about  the  head 
seeming  to  emanate  from  wounds  to  the  mucosa  by 
foreign  bodies,  by  decomposition  of  pieces  of  food  in 
cre^dces  or  by  damage  by  masses  too  large  to  be  swal- 
lowed. In  the  few  instances  where  we  have  tried  bacteri- 
ology, no  definite  result  has  been  obtained  unless  the 
frequent  occurrence  of  organisms  bearing  a  resemblance 
to  Bact.  necrophorus  be  important.  Tliis  organism  how- 
ever may  be  found  in  many  necrotic  processes  in  animals ; 
I  do  not  look  upon  it  as  specific  in  the  locations  just  cited. 

Mammals  as  a  class  do  not  present  many  inflammatory 
conditions  around  the  anterior  head,  aside  from  the 
specific  diseases  like  distemper  (?),  actinomycosis.  Kan- 
garoo disease  and  the  like.  Monkeys  occasionally  have 
acute  coryza,  which  may  indeed  seem  transmissible  to 
others  but  it  seldom  leads  to  any  serious  consequences 
and  is  untreated,  except  by  segregation.  Tuberculous 
lesions  are  not  recorded.  There  has  been  no  glossitis  aside 
from  lesions  involving  the  pharynx.  The  tonsils  have  been 
discussed  under  the  lymphatic  apparatus  and  it  only  need 
be  repeated  here  that  inflammation  and  hypertrophy  of 
these  organs  are  exceedingly  rare.  Specific  or  individual 
diseases  of  the  salivary  glands  are  also  rare  although 
these  organs  may  be  involved  by  extension.  Tliis  gen- 
eral region  is  not  often  affected  with  tumor,  unless  the 
jaw  be  included  which  bone  is  the  seat  of  several 
tumors  in  antelopes  and  opossums.  Aside  from  these 
we  have  seen  an  epithelioma  of  the  tongue  in  a  black  bear 
{TJrsus  americanus) . 

Larynx. 

The  larynx  is  an  organ  of  fairly  uniform  construction 
through  the  mammalian  orders  but  is  conspicuously  dif- 
ferent in  the  Aves  where  it  is  double.  The  upper  end  of 
the  trachea,  in  the  latter  class  is  surmounted  by  a  cartilag- 
inous box  lying  beneath  the  root  of  the  tongue  through 


THE  RESPIRATORY  SYSTEM  139 

which  an  anteroposterior  slit-like  opening  forms  the 
glottis ;  there  is  no  epiglottis.  This  is  only  an  air  passage, 
the  voice  being  made  in  the  syrinx  or  lower  box  which  lies 
at  the  bifurcation  of  the  trachea.  The  structure  of  the 
upper  box  is  quite  simple  with  its  lateral  plates  controlled 
by  the  glossal  muscles  and  two  external  retractors  but 
the  syrinx  is  very  complicated  and  variable  in  the  dif- 
ferent orders  and  even  in  the  same  family.  It  possesses 
an  internal  and  external  set  of  muscles  and  in  some  birds 
can  be  opened  at  one  point  to  permit  air  to  pass  to  the 
cervical  or  thoracic  air  sacs.  Detailed  discussion  of  its 
anatomy  is  hardly  profitable  since  there  is  nothing 
peculiar  about  its  diseases.  On  one  occasion  only  have 
we  seen  distinct  pathological  change — what  was  probably 
an  extension  of  mould  disease  from  it  to  the  cervical  air 
sac.  It  is  involved  in  true  tracheitis  and  bronchitis  but 
even  these  are  rare  in  birds. 

The  larynx  on  the  other  hand  is  constantly  reddened 
in  cases  of  pharyngitis  and  may  be  the  seat  of  mould 
colonies.  Edema  of  this  structure  is,  however,  not  very 
common,  it  being  recorded  but  twice  in  birds  in  associa- 
tion with  nearby  inflammation  and  five  times  in  mammals ; 
in  the  latter  cases  three  were  of  acute  infectious  nature, 
one  was  a  tumor  and  the  other  osteomalacia.  It  is 
common  to  find  the  laryngeal  and  tracheal  mucosa  swollen 
and  wet  in  chronic  bone  degenerations  without  the  con- 
dition being  severe  enough  to  call  it  edema. 

Laryngitis. 

Acute  laryngitis  of  active  catarrhal  or  purulent  nature 
has  been  met  five  times  in  mammals  and  twice  in  birds 
while  more  chronic  lesions  have  occurred  only  in  the 
former,  four  times.  Tuberculous  laryngitis  has  been 
observed  in  a  cockatoo  and  a  lemur;  they  are  interesting 
enough  to  cite.  There  are  no  cases  recorded  among 
monkeys  despite  the  large  number  dying  from  the  disease ; 
this  implies  of  course  that  no  suspicion  of  its  existence 


140  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

was  had  at  postmortem  but  perhaps  some  would  have 
been  detected  had  every  larynx  been  subjected  to  micro- 
scopic section.  A  citron-crested  cockatoo  was  found 
when  posted  to  have  general  miliary  tuberculosis.  The 
bright  red  rim  of  the  glottis  attracted  attention  and  upon 
slitting  open  the  organ,  pinhead  size,  sharply  outlined 
yellow  tubercles  were  found  on  both  sides.  A  black  and 
white  lemur  was  killed  because  of  a  positive  tuberculin 
test.  He  was  in  good  condition  and  exhibited  as  his  only 
lesions  retropharyngeal  lymph  nodes  with  precaseous 
miliary  nodules  and  small  miliary  tubercles  on  the  epi- 
glottis, true  and  false  vocal  chords  and  in  the  mucosa  of 
the  main  ventricle,  each  lesion  being  surrounded  by  a 
narrow  sharply  injected  zone.  This  seems  like  a  recent 
double  implantation  since  the  retropharyngeal  glands 
probably  do  not  drain  toward  or  from  the  larynx.  The 
larynx  has  been  the  seat  of  only  one  tumor,  a  squamous 
cell  cancer  in  an  Azara's  agouti.  The  tumor  caused 
ulceration  and  edema  of  the  whole  mucosa  sufficient  to 
produce  fatal  asphyxia. 

The  trachea  is  of  relatively  little  comparative  or 
pathological  interest  aside  from  its  inflammations  which 
however  are  so  closely  associated  with  bronchitis  that 
they  will  be  included  under  that  heading.  Perhaps  the 
most  important  condition  of  this  tube  is  its  infestation 
with  Syngamus  tracliealis  since  this  leads  to  inflam- 
mations not  only  of  the  related  mucosa  but  predisposes  to 
pulmonary  infection.  The  occurrence  in  the  Galli  is  well 
kno^\Ti  but  perhaps  it  is  not  so  well  recog-nized  that  this 
worm  occurs  also  in  crows  (Passeres)  and  swans  and 
geese  (Anseres).  For  the  diagnosis  of  this  condition  it  is 
customarily  stated  that  a  frothy  mucus  in  the  mouth  is 
very  suggestive;  this  is  true  in  the  cases  seen  here  but 
in  addition  a  mucopurulent  stomatitis  is  exceedingly 
common  and  when  the  two  are  combined  the  picture  is 
almost  confirmatory.  Although  worms  are  credited  with 
considerable  weight  in  the  production  of  pneumonia  in 


THE  RESPIRATORY  SYSTEM  141 

Ungulata,  they  have  only  been  seen   once  within  the 
tracheal  tube. 

The  Beonchi. 

The  bronchi  will  be  discussed  as  a  separate  part  of  the 
respiratory  system  in  so  far  as  possible  since  they  present 
a  very  decided  difference  in  anatomy  between  mammals 
and  birds  and  because  the  incidence  of  their  disease  is 
other  than  will  be  found  for  the  lungs.  However,  distinc- 
tion has  been  made  between  changes  in  the  grosser  tubes 
and  those  in  the  finer  bronchioles,  especially  because 
capillary  bronchitis  so-called  is  really  a  pneumonitis  in 
which  the  mucosa  of  the  larger  passages  need  not  par- 
ticipate. The  mammalian  tubes  are  not  really  greatly  dif- 
ferent in  their  construction,  passing  through  ever  smaller 
branchings  which  give  an  increasing  square  area  of  tube 
capacity  and  more  extensive  mucous  surface.  The  avian 
main  bronchus  breaks  up  very  shortly  after  entering  the 
lung  into  a  varying  number  of  spaces  lined  with  low 
epithelium  lying  upon  a  fibrous  support  and  without 
cartilage.  These  spaces  then  open  into  secondary  air 
spaces  of  a  size  visible  to  the  naked  eye  which  are  in  turn 
surrounded  by  microscopic  alveoli.  The  largest  spaces, 
first  mentioned,  continue  to  grow  smaller  toward  the 
lower  part  of  the  lung  where  they  usually  communicate 
with  one  or  other  of  the  various  ostia  of  air  sacs.  Bron- 
chial diseases  in  birds  must  therefore  be  limited  at  the 
place  where  the  bronchi  lose  the  cartilaginous  rings  since 
below  this  the  surface  functionates  as  pulmonary  tissue. 

Beonciiitis. 

The  accompanying  list.  Table  8,  will  show  the  distri- 
bution of  bronchitis  not  accompanying  pneumonia  or  due 
to  mycosis.  It  is  striking  that  carnivorous  animals  are 
more  prone  to  bronchitis  than  any  other  order  (the 
struthious  birds  are  too  few  to  be  important).  There  is  a 
very  decided  preponderance  of  mammalian  cases  over 
a\4an,  there  being  not  only  more  cases  but  proportion- 


142   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 


ately  more  orders  affected.  The  character  of  lesions  in  the 
mammals  is  nearly  always  catarrhopurulent  or  freely 
purulent  while  ulcerative  changes  are  not  uncommon. 
Peribronchial  infiltrates  are  seldom  found  without  some 

Table  8. 

Showing  the  Percentage  Incidence  of  Bronchitis  and  of  Parasites  in  the  Autopsies 
upon  the  Various  Orders. 


Orders 

Simple  Bronchitis 

Verminous  Bronchitis 
and  Pneumonitis 

Inactive  Parasites  of 
Lung  (Encysted)  &c. 

Cases 

Per  cent. 

Cases 

Per  cent. 

Cases 

Per  cent. 

Primates 

7 
14 

5 

3 

4 

4 

1 

1 

2 
1 

1.4 
2.9 

1.3 

1.7 

.29 

.58 
.5 

1. 

.67 
3.3 

3 

8 

2 
2 

2 

7 

1 
3 

.6 
1.6 

1. 
.5 

1.1 
.5 

.3 
1. 

5 

1 
8 

10 

2 
3 

2 

1 

1. 

Lemures 

Carnivora 

Insectivora 

Chiroptera 

Rodentia 

Ungulata 

Proboscidea 

Hyracoidea 

Edentata 

1.2 
1.6 

2.7 
12.5 

Marsupialia 

Monotremata 

Passeres  . 

1.7 
.16 

Picariae 

Psittaci 

Accipitres 

.GaUi 

Hemipodii 

Fulicariae. 

Alectorides 

Limicolse 

Gaviae 

Steganopodes 

Herodiones 

Odontoglossse  .... 

Palamedes 

Anseres 

1. 

Struthiones 

For  meaning  of  italics  see  footnote  Table  1. 

evidence  of  pneumonia ;  nor  is  it  common  to  meet  the  pale 
lines  extending  from  bronchi  between  the  lobules,  such  as 
are  seen  in  human  streptococcal  disease.  Avian  bron- 
chitis is  usually  hemorrhagic  or  catarrhal  and  with 
exceeding  rarity  becoming  purulent ;  when  this  occurs  the 


THE  RESPIRATORY  SYSTEM  143 

cause  is  frequently  found  to  be  tuberculosis  or  mycosis. 
Inflammation  of  the  larger  passages  is  nearly  always 
accompanied  by  pulmonary  congestion, ,  a  serious  con- 
dition in  birds  as  will  be  seen  later. '  A  few  of  these  cases 
have  been  studied  bacteriologically  with  no  definite  result, 
nor  have  these  cases  occurred  in  such  groups  that  an 
epizootic  was  suggested.  Bact.  avisepticum,  Bact.  cani- 
septicum,  Bact.  coli,  Bact.  aerogenes  mucosum,  and  Ps. 
pyocyaneus,  Streptococcus  hemolyticus  and  non-hemo- 
lyticus  and  staphylococci  have  been  found. 

A  review  of  the  active  verminous  lesions  of  the  bronchi 
and  the  pneumonitis  to  which  parasites  lead,  shows  again 
the  liighest  number  among  the  Carnivora,  with  negligible 
percentages  among  the  Aves.  The  forms  concerned  are, 
when  determined,  ascarides,  strongylus,  hepaticola,  fas- 
ciolopsis,  cytoleichus,  pneumonyssus  and  paragonimus; 
these  will  be  discussed  later.  The  lesions  in  the  bronchi 
are  mucocatarrhal  or  hypertrophic;  occasionally  actual 
ulcerations  are  seen.  What  is  more  important  however  is 
the  peribroncliitis  leading  to  interstitial  pneumonitis  and 
to  bronchiectasis  of  the  smaller  bronchi,  or  to  areas  of 
atelectasis  by  total  occlusion  of  some  small  air  passage. 
This  pathology  is  fairly  well  recognized  among  veterinary 
pathologists  but  there  are  two  points  which  seem  worthy 
of  special  emphasis,  namely,  the  relative  mildness  of  the 
changes  in  the  larger  bronchi  and  the  importance  of  the 
worms  as  causes  of  pneumonia.  In  regard  to  the  first 
it  can  be  pointed  out  that  the  trachea  and  its  branches 
need  not  be  altered  at  all  while  the  middle  sized  bronchi 
present  a  mottling  of  small  recent  congestion  with  pig- 
mentations from  old  hemorrhages,  together  with  slight 
unevennesses  of  the  surface.  Small  bronchi  on  the  other 
hand  are  the  seat  of  ulcerative  internal  processes  and 
quite  marked  peribronchitis,  as  indicated  by  round  and 
connected  tissue  nuclei  or  perhaps  polynuclear  increase 
under  active  acute  inflammation;  it  is  in  the  latter  case 
that  acute  pneumonitis  is  present. 


144  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

To  what  extent  do  the  parasites  predispose  to  pneu- 
monia? Compare  for  this  purpose  the  two  columns  of 
Table  8  showing  active  and  inactive  pulmonary  parasi- 
tism. In  Rodentia  and  Galli  alone  do  we  note  that  active 
parasitism  is  effective,  there  being  no  passive  cases.  In 
all  the  other  orders,  animal  invaders  of  the  lung  are  more 
often  encountered  as  quiescent  or  encapsulated  bodies, 
therefore  as  findings  incidental  to  the  autopsy  and  per- 
haps not  concerned  in  the  cause  of  death.  In  Ungulata 
the  inactive  parasitisms  are  five  times  as  frequent  as  the 
active  inflammatory  lesions.  It  might  be  added  that  the 
list  is  made  up  of  cases  wherein  we  saw  parasites  whether 
determined  or  not,  an  explanation  of  the  apparently  small 
number  of  cases;  there  were  many  more  in  which  such 
invaders  were  suspected  but  not  found  and  there- 
fore excluded. 

Beonchiectasis. 

Bronchiectasis  affecting  the  smallest  tubes,  or  bron- 
chiolectasis,  is  not  at  all  uncommon  in  verminous  pneu- 
monitis and  is  explained  as  due  to  the  degeneration  of  the 
wall,  the  surrounding  progressive  ulceration,  to  accumu- 
lation of  inspired  air  and  its  retention  by  the  obstruction. 
There  is  described  a  generalized  bronchiolectasis,  chiefly 
in  young  human  beings,  due  to  a  destructive  bronchio- 
litis ;  this  has  not  been  seen. 

Non- verminous  bronchiectasis  of  the  middle  sized 
bronchi  such  as  is  seen  in  human  chronic  bronchitis, 
simple  or  tuberculous,  is  quite  uncommon.  Widening  of 
the  bronchial  lumen  may  be  divided,  as  I  see  its  path- 
ogenesis, into  (a)  that  due  to  congenital  weakness  of  the 
walls,  (b)  that  due  to  obstruction  permitting  air  to  pass 
into  but  not  out  of  a  bronchus  because  of  a  ball-valve 
obstruction  or  weakness  of  expiratory  power,  (c)  that  due 
to  external  pressure  by  tumors  or  distortion  by  fibrous 
tissue  either  witliin  the  lung  or  pleura  and  (d)  that  due  to 
inflammatory  weakening  of  walls,  augmented  by  loss  of 


THE  RESPIRATORY  SYSTEM  145 

supporting  pulmonary  tension,  accummulation  of  secre- 
tion and  the  dilating  etfect  of  inspiration  preparatory  to 
and  incidental  to  coughing.  How  important  the  last 
three  auxiliary  factors  may  be  in  the  cases  explicable 
under  a,  b,  c,  can  easily  be  speculated  upon  and  may  vary 
in  different  cases. 

Bronchiectasis  is  reasonably  common  with  pulmonary 
diseases  of  man,  particularly  of  chronic  character,  but 
is  certainly  not  frequent  among  animals.  Under  the  first 
group  (a)  we  can  record  one  case,  a  Siberian  tiger  which 
died  of  enteritis  and  its  complications  to  which  were  added 
a  mild  inactive  bronchitis  and  a  broncliiectasis  of  diffuse 
distribution.  The  lungs  were  irregular  in  shape,  dull, 
gray  red  in  color  and  gave  a  variable  boggy  and  vesicular 
sensation  to  the  fingers.  On  opening  the  lung,  dilatations 
of  the  bronchi  were  found,  affecting  chiefly  the  larger 
secondaries  but  apparently  not  the  broncliioles.  Para- 
sites were  not  found  nor  were  inflammatory  reactions 
apparently  adequate  to  explain  the  distentions,  so  that  we 
looked  upon  this  case  as  congenital.  Cases  coming  under 
the  headings  b  and  c  are  not  recorded.  Inflammation 
almost  certainly  represents  the  most  important  single 
factor  in  the  pathogenesis  of  this  lesion  and  could  be 
demonstrated  in  two  cases,  a  Clouded  Leopard  {Felis 
nehulosa)  and  a  Red  River  Hog  {Potmnockcerus  porcus). 
While  I  feel  that  parasites  probably  laid  the  foundation 
for  the  dilatations  in  these  cases,  none  were  found  after, 
in  one  case  at  least,  a  very  thorough  search,  although  in 
the  second  animal  a  single  cyst  of  Cysticercus  tenuicollis 
was  found  in  the  peritoneum.  In  both  animals  there  was 
a  low  grade  interstitial  pneumonitis  and  peribronchitis 
mth  dilatations  of  the  middle  sized  and  end  bronchi,  these 
being  supplied  mth  thick  walls  but  containing  very 
scanty  secretion. 

We  have  on  record  chronic  ulcerative  pulmonary 
tuberculosis  in  six  primates,  two  carnivores  and  nine 
ungulates.     It  is  highly  probable  that  among  this  number 


146   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

some  cases  of  ulcerative  bronchiectasis  occurred  but  if  so 
they  were  not  conspicuous  enough  to  mention  in  the  diag- 
nosis and  in  only  two  protocols  do  I  find  a  discussion 
thereof,  once  in  a  monkey  and  once  in  a  carnivore.  All  the 
cases  of  our  records  were  diffuse  ectasias,  fusiform,  or 
irregular  and  none  of  the  distinct  saccular  variety. 

From  the  foregoing  facts  it  would  seem  that  in  human 
cases  more  weight,  should  be  laid  to  the  effect  of  the 
dilating  power  of  coughing  and  its  preparatory  exertions. 
While  I  am  aware  that  the  comparative  incidence  of 
human  and  lower  animal'  bronchiectasis  cannot  be  based 
upon  the  meager  figaires  at  our  command,  these  dilata- 
tions certainly  can  be  expected  in  a  general  autopsy 
ser\dce  more  in  man  than  in  lower  animals.  Chronic 
bronchitis  is  relatively  rare,  aside  from  the  verminous 
varieties.  I  have  seen  little  retained  exudate  in  the 
bronchi,  probably  because  quadrupeds  seem  with  ease  to 
raise  and  swallow  the  secretions.  Nor  do  animals  give 
vent  to  paroxysms  of  coughing  such  as  the  human  being 
feels  forced  to  do.  Suggestive  deductions  from  these 
points  are  that  inflammation  is  the  principal  factor  in 
acquired  bronchiectasis  and  that  the  retention  of  secretion 
with  violent  inspiratory  efforts  are  potent  in  man  for  the 
dilatation  of  the  tubes. 

The  Lungs. 

The  essential  respiratory  organ  of  the  animal  body, 
the  lung,  is  all  through  this  kingdom  a  structure  intended 
to  expose  the  blood  to  free  or  combined  atmosphere  in 
order  to  permit  gaseous  interchange,  therefore  being 
arranged  so  that  there  is  a  close  apposition  of  the  two 
factors,  separated  only  by  such  cells  and  membranes  as 
may  be  necessary  to  protect  the  circulation ;  perhaps  these 
anatomical  elements  possess  at  the  same  time  some  vital 
force  to  further  the  exchange  of  useful  and  useless  matter. 
In  the  two  classes  under  discussion  there  is  no  difference 
whatsoever  in  the  primary  factors  of  respiration  albeit 


THE  RESPIRATORY  SYSTEM  147 

some  variations  exist  in  reference  to  oxygen  and  carbon 
dioxide  interchange,  moisture  of  the  air,  and  the  physics 
of  inspiration  and  expiration.  The  chemical  variables 
have  in  our  limited  knowledge  of  comparative  physiology 
apparently  little  effect  upon  morbid  anatomy  but  it  is 
probable  that  some  pathology  may  be  in  part  explained 
on  physical  grounds. 

The  mammalian  respiratory  box  is  a  relatively  elastic 
affair,  but  collapsed  at  the  end  of  expiration  which  is 
largely  a  passive  or  recoil  process,  f  The  avian  thorax  is 
believed  to  be  normally  a  tensely  distended  space  from 
which  air  is  expelled  by  pressure  of  the  pectoral  contrac- 
tion upon  the  broad  sternum  driving  the  latter  back  upon 
the  air  sacs  which  in  turn  drives  it  from  the  lungs.  Also 
by  this  means,  air  is  distributed  through  the  bones  and- 
air  spaces,  a  measure  necessary  in  flight,  especially  in  a 
head  wind  when  tracheal  inspiration  is  said  to  be  sus- 
pended at  times..  The  communications  of  the  lungs,  air 
sacs  and  bones  make  it  possible  for  birds  to  breathe 
internally  when  the  trachea  is  closed  and  externally  as 
well  if  a  bone  be  opened.  The  balance  of  air  pressure  in 
the  lungs  and  related  spaces  is  dependent  upon  the 
patency  of  the  ostia  communicating  between  the  bronchial 
ends  and  the  air  sacs,  from  wliich  the  bony  cavities  obtain 
their  supply.  Should  all  these  be  closed  there  is  first  a 
standstill  of  current  and  a  limitation  of  the  respiration  of 
the  lungs.  Fortunately  it  is  extremely  rare  that  this 
occurs  for  it  is  obvious  that  it  is  incompatible  with  flight, 
and  with  life  indeed.  The  principal  effect  upon  the  lungs 
of  obstruction  to  the  passages  seems  to  be  expressed  in 
congestion  but  in  how  far  this  is  due  actually  to  the 
closure  of  foramina  and  how  far  to  the  cause  of  obstruc- 
tion is  sometimes  difficult  to  evaluate.  '  It  should  be 
remembered  that  the  air  sacs  are  usually  looked  upon  as 
mucous  surfaces  continuous  with  the  bronchial  wall,  there 
being  a  deep  layer  to  each  membrane  possibly  continuous 
with  the  serous  membranes.     In  mould  disease  of  the 


148  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

lungs  there  is  very  commonly  a  colony  lying  in  the  ostium 
supplying  the  anterior,  lateral  and  posterolateral  ca\"ities.^| 
1  The  lungs  in  birds  are  not  free  as  in  mammals,  being 
fitted  into  the  troughs  made  by  the  anterior  ridges  of  the 
ribs,  to  the  serous  covering  of  which  thej^  are  lightly 
attached  by  delicate  fibres  running  between  the  two.  This 
more  or  less  definite  fixation,  together  with  the  pressure  of 
the  air  in  the  sacs  give  the  free  play  of  the  lungs  a  limited 
excursion.  They  are  naturally  very  elastic  by  reason  of 
a  good  supply  of  elastic  fibers  and  large  air  spaces,  a  con- 
dition aided  by  their  attachments  to  the  supports  of  the 
diaphragm  and  to  the  insertions  of  the  air  sac  walls.  Not- 
withstanding this  elasticity  and  the  great  capacity  of  the 
organ  for  blood,  it  seems  as  if  congestion  of  the  lungs  is 
a  very  serious  matter,  since  from  the  foregoing  re^dew 
of  anatomy,  accommodation  of  excess  blood  and  any  con- 
solidation must  be  difficult.  As  a  matter  of  fact  the  mere 
excess  of  blood  known  as  active  congestion  seems  able  to 
kill  small  varieties.J 

Congestion-  op  Lungs. 

Birds  of  flight  seem  to  have  little  resistance  to  this  con- 
dition and  often  it  is  the  only  diagnosis  one  can  make  at 
autopsy.  The  causes  of  this  condition  include  exposure, 
dust,  gorging  (?),  indigestion,  enteritis  and  infection  in 
birds  while  in  mammals  acute  gastrointestinal  disease 
stands  out  as  the  most  prominent  accompaniment.  To 
what  extent  dust  and  exposure  operate  I  do  not  see, 
although  they  are  frequently  mentioned  as  causes.  The 
overfilling  of  the  crop,  esophagus  and  proventricle,  the 
turgescence  incident  to  gastric  indigestion  or  the  pressure 
of  foreign  bodies  in  large  amount  are  supposed  to  operate 
by  exerting  pressure  on  the  anterolateral  air  sacs  mth 
closure  of  their  ostia  and  also  by  right  lateral  torsion  of 
the  heart  with  twisting  of  the  very  delicate  pulmo- 
nary veins.  , 


THE  RESPIRATORY  SYSTEM  149 

I  have  sought  to  show  that  protozoa  or  embryo 
nematodes  in  the  blood  might  embarrass  the  lungs  to  a 
state  of  congestion,  a  thought  suggested  by  some  findings 
in  the  London  Gardens,  but  only  about  ten  per  cent,  of  our 
cases  of  hemic  parasitism  are  accompanied  by  it. 

The  incidence  of  congestion  of  the  lungs  not  due  to 
stasis  as  from  cardiac  diseases,  is  2.4  per  cent,  in  mam- 
malian autopsies,  in  only  7  per  cent,  of  which  figure  did  it 
represent  the  principal  morbid  anatomy,  whereas  in  birds 
it  occurred  to  the  extent  of  7.6  per  cent,  of  postmortems, 
in  17  per  cent,  of  which  it  was  the  sole  or  principal  cause 
of  death.  This  seems  to  bear  out  the  feature  of  delicacy 
of  the  pulmonary  vascular  mechanism  in  these  latter 
animals.  This  condition  seems  to  be  indicated  by  simple 
dyspnoea  in  birds,  relief  for  which  has  occasionally  been 
afforded  by  removal  from  the  exhibition  cages  and  pro- 
tection separately  in  a  warm  dry  room;  this  is  partly 
hypothetical  of  course  and  congestion  is  to  be  looked  upon 
as  serious,  particularly  in  passerine  birds. 

Pneumonia. 

Pneumonia  as  a  clinical  disease  is  a  relatively  un- 
common, although  quite  serious  sporadic  condition  in 
animals.  However  accompanying  the  specific,  more  or 
less  epizootic  diseases  such  as  influenza,  distemper  and 
the  choleras  it  may  be  a  frequent  and  quite  pronounced 
complicating  feature  of  the  case.  Pneumonia  per  se  has 
exacted  a  reasonable  toll  in  this  Garden  but  unfortunately 
recognition  being  impracticable,  diagnosis  and  treatment 
have  not  progressed.  Nor  has  it  been  practicable  to  group 
our  cases  pathologically  because  of  the  lack  of  history 
and  the  difificulty  of  making  bacteriological  observations 
at  many  autopsies.  Fortunately  we  have  had  practically 
no  epizootic  pneumonias,  an  experience  shared  with  other 
gardens  judging  by  their  published  reports.  Etiologi- 
cally,  and  of  course  this  applies  to  non-verminous,  non- 
mycotic  and  non-tuberculous  cases,  the  pneumococcus  has 


150   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 


stood  out  prominently  as  a  cause  with  a  few  additional 
cases  due  to  the  streptococcus  and  the  Bact.  aerogenes 
mucosum  group ;  London  reports  four  cases  in  monkeys 
due  to  the  Friedlander  bacillus.     Some  time  ago  Doctor 

Table  9. 

Showing  the  Number  of  Cases  of  the  Various  Forms  of  Pneumonia   Found  in 
Each  of  the  Orders. 


Primates 

Lemures 

Carnivora. . .  . 
Insectivora. . . 
Chiroptera .  .  . 

Rodentia 

Ungulata .  .  .  . 
Proboscidea .  . 
Hyracoidea  .  . 
Edentata .  .  .  . 
Marsupialia.  . 
Monotremata 

Passeres 

Picariae 

Striges 

Psittaci 

Accipitres. .  .  . 
Columbaj .  .  .  . 

GalU 

Hemopodii.  .  . 
Fulicariae  .  .  .  . 
Alectorides. . . 

Limicolse 

Gaviae 

Impennes.  .  .  . 
Steganopodes. 
Herodiones. . . 
Odontoglossse. 
Palamedes .  .  . 

Anseres 

Struthiones .  . 


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Weidman  subjected  our  pneumonias  to  an  analysis  and 
was  able  to  show  that  there  is  no  parallelism  between  the 
seasonal  incidence  of  pneumonia  in  man  and  animals, 
rather  indeed  that  the  Garden  is  more  apt  to  have  a 
greater  number  of  cases  in  the  summer,  a  sort  of  *'  closed 


THE  RESPIRATORY  SYSTEM  151 

season  ' '  for  man.  This  I  am  inclined  to  interpret  as  con- 
nected with  the  larger  number  of  visitors  during  that 
season.  Doctor  Weidman  was  further  able  to  show  that 
the  only  real  examples  of  lobar  fibrinous  pneumonia 
strictly  comparable  to  the  human  infection  occurred  in  the 
Primates.  I  have  uncovered  one  in  a  lemur  and  one  in  a 
carnivore.  The  pneumococcus  has  been  far  and  away  the 
greatest  producer  of  our  pneumonias,  in  two  typed  cases 
being  of  the  IV  variety.  There  mil  be  given  below  a 
summary  of  the  pathological  types  of  pneumonia  en- 
countered, to  be  followed  by  some  notes  upon  the  prin- 
cipal gross  and  minute  anatomy  in  special  orders.  Table 
9  mil  show  the  numerical  distribution  of  types  among 
the  orders.  All  the  principal  mammalian  orders  are 
represented  while  the  birds  seem  relatively  less  sus- 
ceptible to  the  disease  and,  except  the  Passeres,  show  a 
trifling  incidence. 

Primates  present  a  definite  group  of  variations  from 
the  other  orders,  notably  in  having  four  clear  cases  of 
lobar  fibrinous  pneumonia,  and  in  certain  histological 
findings.  In  reference  to  the  lobar  cases,  a  re\aew  of  their 
history  does  not  indicate  that  any  might  have  been  surely 
diagnosed  by  their  symptoms,  and  only  possibly  by  signs 
in  one  case  during  the  stage  of  red  hepatization;  unfortu- 
nately no  temperature  records  are  at  hand.  In  one  case 
it  was  possible  to  see  a  group  of  alveoli  wdth  the  fibrin 
collected  in  a  strand  which,  according  to  classical  descrip- 
tion, passes  through  the  septum  to  the  adjoining  alveolus. 

There  were  two  cases,  a  Chimpanzee  (Pan  niger)  and 
a  Galago  (Gal ago  maholi)  with  a  microscopical  picture 
suggestive  of  those  we  met  in  the  influenza  epidemic,  and 
indeed  the  lung  of  the  former  resembles  grossly  the  lung 
of  influenza  pneumonia.  The  spotty  areas  of  watery 
purple  color  correspond  under  magnification  to  cellulo- 
edematous  semisolid  sections  shomng  a  sanguineous 
exudate,  fewpolynuclear  cells  and  many  swollen  epithelia. 
The  microscopic  picture  of  the  bronchocatarrhal  pneu- 


152  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

monias  shows  conspicuously  thickened  septa  decidedly 
^\dder  than  one  is  accustomed  to  see  in  human  cases  and 
apparently  due  more  to  round  cell  infiltration  than  to  con- 
gestion or  polynuclear  increase. 

Bronchopneumonia  or  capillary  broncliitis  with  zones 
of  cellular  edema  in  the  vicinity  is  a  rather  usual  picture 
in  the  deaths  from  degenerative  bone  disease.  It  cannot 
be  said  that  there  is  anything  very  peculiar  about  it, 
although  a  frequent  note  met  in  the  autopsies  describes 
spotty  areas  of  hemorrhage  and  nearby  atelectasis. 

The  case  of  lobar  pneumonia  found  in  a  ring-tailed 
lemur  {Lemur  catta)  showed  very  delicate  fibrinous  retic- 
ulum and  relatively  few  cells  in  the  exudate,  a  picture 
apparently  due  in  part  to  beginning  resolution  since  the 
whole  upper  left  lobe  was  in  a  stage  of  gray  hepatization. 

The  peculiarity  of  the  Carnivora  seems  to  lie  in  the 
reaction  of  the  epithelia,  these  cells  being  quite  large, 
swollen  and  occasionally  much  vacuolated.  Such  a  picture 
was  most  pronounced  in  the  terminal  bronchitic  pneu- 
monias in  cases  which  might  be  called  distemper.  Many 
instances  of  pseudolobar  catarrhal  or  bronchopneumonia 
are  recorded  but  we  also  observed  the  fibrinous  lobar 
form  at  the  stage  of  red  hepatization  in  a  Texas  skunk 
{Mephitis  mesomelas) .  Concerning  the  orders  Rodentia 
and  Edentata  no  especial  notes  seem  necessary  for  their 
inflammatory  reactions  are  essentially  like  the  others  in 
that  epithelial  cells  are  much  swollen  and  prominent. 

Pneumonias  of  Ungulata  are  well  known  to  pathology 
and  offer  in  causation  and  microscopy  little  that  is 
peculiar.  It  might  be  emphasized  however  that  the  gross 
appearance  of  the  bronchocatarrhal  variety  closely  simu- 
lates that  of  lobar  pneumonia,  therefore  to  be  called  a 
pseudolobar  form,  in  that  extension  to  various  parts  of 
a  lobe  seems  to  occur.  Moreover  in  the  bronchitic 
varieties  associated  with  enteritis,  with  or  without  infec- 
tious foci  in  the  pharynx  or  larynx,  there  may  be  two  or 
even  three  stages  of  the  pneumonitic  process  in  one  lung 


hl(      8— \()R\I\I      WIW     IKI\l\k"l      \\i)    -UONDXk'i     \l  \  1  Ol  I 
THi      M  n  AI      PROLONG  M  lON-i    IHAI     1  (H  M)      IHl      J'RI\IAR\       \1  \  1  i 


Fig.  9. — EARLY  HRONCHOPNKUMONIA  OF  SUPERFICIAL  ORIGIN.  NOTE  SOME  LITTLE 
EXUDATE  IN  SECONDARY  ALVEOLUS.  WIDE  SWOLLEN  SEPTA  AND  BOTH  ALVEOLI  REDUCED 
IN   SIZE. 


THE  RESPIRATORY  SYSTEM  153 

or  lobe.  It  seems  that  this  pseudolobar  appearance 
occurs  definitely  more  often  in  ungulates  than  in  the 
other  orders. 

Marsupialia  offer  two  rather  easily  grouped  classes 
of  bronchopneumonia — one  associated  with  enteritis  and 
one  secondary  to  "  Kangaroo  disease  "  of  the  jaw;  they 
differ  in  microscopy  correspondingly.  The  simple  bron- 
chitic  and  peribrOnchitic  infiltrate  and  superficial  exudate 
occurring  with  enteritis  or  with  a  general  infection  is 
relatively  diffuse,  giving  in  some  instances  the  impression 
of  an  interstitial  process  and  showing  notably  swollen 
septa;  there  may  be  fibrin  but  this  is  exceptional  and 
scanty.  Wlien  mycosis  of  the  jaw  has  been  the  origin  or 
occasion  of  the  infection  the  picture  is  that  of  frank  aspi- 
ration pneumonia,  therefore  more  like  a  septic  infarct. 
However  the  amount  of  fibrin  is  sometimes  very  great  and 
whole  alveoli  will  be  filled  with  it,  perhaps  accompanied 
by  red  cells,  polynuclears  and  epithelia.  Epithelial  cells 
however  play  a  small  part  in  the  minute  anatomy.  Hemor- 
rhage and  edema  are  prominent  but  true  abscess  forma- 
tion and  gangrene  are  not.  Possibly  the  animals  die  too 
soon  for  the  latter  to  develop. 

Pneumonia  in  Aves  aside  from  that  due  to  moulds  is 
apparently  much  less  coromon  than  among  the  Mammalia, 
one  order  only,  the  Passeres,  showing  an  incidence  com- 
parable to  the  important  orders  of  the  latter  class.  The 
other  orders,  and  this  applies  particularly  to  those  of 
which  we  have  an  adequate  number,  are  quite  insus- 
ceptible to  simple  pneumonia,  none  of  them  showing  over 
two  per  cent.  /  There  are  listed  for  Aves  three  instances 
of  lobar  fibrinous  pneumonia.  These  cases  can  be 
described  together  since  in  all  the  findings  were  about  the 
same.  A  whole  lung  or  goodly  portion  thereof  was  uni- 
formly involved  in  a  red  or  gray  consolidation  of  rather 
fine  granular  character  which  on  section  study  seemed  to 
be  made  up  of  the  same  lesion  all  over,  with  fibrin  a  promi- 
nent part  of  the  exudate.  The  coagula  were  largely 
11 


154  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

within  the  secondary  alveoli  but  the  primaries  also 
contained  it.  The  microscopic  section  may  not  have  rep- 
resented the  process  at  all  places,  and  since  the  arrange- 
ment of  fibrin  is  similar  in  definitely  catarrhal  lesions, 
these  may  of  course  have  been  instances  of  pseudo- 
lobar  pneumonia. 

Our  data  are  too  few  to  draw  any  conclusions  as  to  the 
behavior  of  the  various  orders  but  one  note  may  be  per- 
mitted. The  passerine  birds  have  a  great  tendency  to 
dense  cellular  infiltrates  while  parrots  show  more  coagu- 
lative  or  fluid  exudates. 

Production  of  Insular  Pneumonia  in  Birds. 

Insular  consolidations  in  which  catarrhal  and  infiltra- 
tive processes  are  prominent,  the  bronchopneumonias, 
seem  to  arise  in  two  ways.  One  course  of  events  appar- 
ently follows  infection  via  the  bronchial  mucosa,  the  other 
via  the  blood  stream  and  a  study  of  the  resulting  lesions 
may  help  toward  an  understanding  of  the  development  of 
pneumonia  in  man. 

When  infection  unquestionably  has  been  superficial, 
that  is  via  the  bronchus,  the  first  thing  to  happen  is  a 
swelling  of  the  septal  prolongations  dividing  the  primary 
alveoli  and  an  extension  of  their  ends  farther  into  the 
secondary  alveoli  with  the  result  that  the  inlet  to  the 
primary  air  sacs  is  narrowed  and  the  space  in  the  second- 
aries is  reduced.  Upon  the  surfaces  there  then  develops 
the  usual  catarrhal  exudate  while  in  the  deeper  parts 
marked  congestion  makes  its  appearance.  Fibrin  may 
develop  and  be  mixed  with  the  cells  both  in  the  larger  and 
smaller  alveoli  but  it  is  more  e\ddent  in  the  former. 
(Figs.  8,  9, 10.) 

The  other  process  by  which  insular  pneumonia  de- 
velops seems  to  begin  in  the  septa  of  the  smaller  alveoli 
and  in  the  perivascular  areas.  This  has  been  looked  upon 
as  hematogenic  or  pleurogenic.  The  first  change  occurs  in 
the  surroundings   of  the  primary  alveoli  where  there     ) 


Fig.  10. — LATER  BRONCHOPNEUMONIA  OK  SUPERFICIAL  ORK;iN.  I 
GREATER  SWELLING  OF  SEPTA.  PRIMARY  ALVEOLI  PRACTICALLY  ALL 
HAS  BECOME  CONSOLIDATED. 


Fir..  11. — INSL'I.AR  PNiaMUMA,  Bl.t,lNMNC,  A>  Cl.l.lA  LAK.  IMILIRAIION  OF  DEHl'ER 
PARTS  OF  SEPTA  AND  OF  INTERSTITIAL  TISSLE.  FOUR  AREAS  OF  DENSE  AIRLESS  CONSOI.ID.A- 
TION.     ALL  SECONDARY  AND  .MANY  PRIMARY  ALVEOLI  WIDELY  OPEN. 


THE  RESPIRATORY  SYSTEM  155 

appears  a  richness  of  nuclei,  of  round,  moderately  well 
stained  character,  among  which  one  may  see  a  few  gran- 
ular and  red  blood  cells.  Soon  the  epithelia  of  adjacent 
alveoli  increase  in  number  and  a  fibrinocellular  exudate 
appears,  at  first  probably  in  the  smaller  sacs.  However 
when  the  lesion  is  intensive  the  course  of  events  must  be 
rapid  for  the  identity  of  a  group  of  primary  alveoli  is  soon 
lost  and  the  exudate  may  extend  to  the  larger  air  space. 
(Fig.  11)  In  severe  or  late  cases  a  decision  as  to  the 
course  of  origin  is  often  impossible.  The  most  instructive 
point  of  this  part  of  the  study  is  the  closing  of  primary 
alveoli  by  the  sw^elling  of  their  septal  ends  and  the  early 
occlusion  of  the  secondary  alveolus  by  a  catarrho- 
fibrinous  or  even  pus-like  material.  It  is  quite  possible 
that  a  similar  course  of  events  transpires  in  the  path- 
ogenesis of  human  pneumonia,  the  superficial  avian  form 
being  comparable  to  the  aspiration  form,  the  interstitial 
form  comparable  to  the  septicemic  variety.] 

Abscess  and  Gangrene  of  Lung. 

Abscess  and  gangrene  of  the  lung  are  degenerative 
processes  dependent  upon  embolism,  or  inspiration  of 
infective  matter  and  it  is  usually  assumed  that  gangrene 
succeeds  upon  abscess  when  the  blood  or  air  supply  of  a 
part  of  the  pulmonary  tissue  has  been  obstructed 
mechanically  or  by  inflammation.  A  review  of  our 
material  adds  little  to  the  etiology  or  pathogenesis  of 
these  two  lesions,  well  recognized  as  they  are  by  veterina- 
rians. As  opposed  to  human  beings,  lower  animals  prob- 
ably suffer  more  from  them,  for  an  explanation  of  which 
one  can  probably  look  to  the  B.  necrosis  or  necrophorus, 
an  organism  quite  common  in  feed,  and  acknowledged  to 
be  of  great  importance  as  a  secondary  invader  during 
specific  infectious  diseases.  It  has  been  found  in  embolic 
abscesses  and  in  the  organs  in  calf  diphtheria  and  similar 
other  conditions.  It  has  been  cultivated  here  twice,  once 
from  a  lung  abscess,  once  from  Kangaroo  disease.     It 


156  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 


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158  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

doubtless  occurs  in  human  necrotizing  processes  but  is 
seldom  emphasized  or  even  heard  about;  possibly  none  is 
due  to  it  or  its  congeners. 

The  distribution  of  abscess  and  gangrene  in  the  lungs 
in  terms  of  the  antecedent  disease,  therefore  its  causation, 
may  however  be  of  interest.  The  lower  animals  move 
more  in  the  horizontal  position,  they  seldom  cough,  they 
are  subject  to  several  different  diseases  A\T[th  principal 
lesions  in  the  anterior  head  (diphtheria,  actinomycosis, 
etc.)  but  not  to  chronic  lymphatic  infection,  they  push 
their  snouts  into  all  kinds  of  filth  thereby  probably  taking 
into  the  nose  and  throat  many  objects  which  can  find  their 
way  to  the  bronchi,  and  finally  they  are  not  subjected  to 
various  instrumental  operative  procedures  when  they 
chance  to  have  a  focus  of  pathology  in  the  nasopharynx. 
For  these  reasons  the  position  of  abscess  and  its  sequels 
may  be  instructive.  It  has  recently  been  stated  that  ab- 
scesses of  the  lung  in  human  beings  following  anesthesia 
for  infections  of  the  upper  respiratory  tract,  occur  most 
often  in  the  upper  parts  of  the  lung  whereas  those  follow- 
ing pneumonia  develop  more  in  the  lower  lobes.  For  the 
animals  of  this  series,  these  facts  are  not  borne  out.  In 
the  accompanying  list  will  be  found  our  acceptable  cases 
of  abscess  and  gangrene.  Little  can  be  said  about 
incidence  save  the  fact  that  the  highest  percentage  and 
number  occur  in  the  Carnivora.  It  will  be  found  that 
the  right  lung  is  affected  nine  times  predominatingly 
while  the  left  lung  is  affected  seven  times.  The  lobe  most 
often  singled  out  for  an  isolated  lesion  is  the  lower  left, 
the  right  middle  being  the  next  most  often  affected.  These 
figures  concern  the  mammals  alone,  the  three  birds  being 
considered  too  few  to  discuss.  It  cannot  be  said  from 
these  figures  that  there  is  in  animals  any  definite  distri- 
bution of  pulmonary  abscess  and  gangrene. 

Nor  do  these  figures  correspond  to  those  appearing  in 
literature  of  human  pathology.  In  man  inspiration  of 
foreign  bodies,  including  bacteria-laden  mucus,  usually 


THE  RESPIRATORY  SYSTEM  159 

carries  them  into  the  right  lung  because  of  the  larger 
and  more  directly  vertical  bronchus  to  that  side.  Emboli 
go  as  a  rule  also  to  the  right  lung  more  than  to  the  left 
because  of  the  greater  size  of  the  pulmonary  artery  and 
more  direct  blood  current  to  that  side.  In  these  animals 
right  side  lesions  are  more  numerous  but  the  left  lower 
lobe  is  the  principal  segment  of  the  lung  to  be  affected. 
The  cases  are  too  few  and  the  anatomy  too  variable  to 
permit  any  deductions.  There  is  in  the  affected  animals 
no  uniform  anatomical  peculiarity  which  would  explain 
the  predominance  of  the  right  lung  as  a  whole  or  the  left 
lower  lobe  as  a  unit. 

Pneumonokoniosis. 

Pneumonokoniosis,  because  of  its  importance  in  indus- 
trial diseases,  has  been  subjected  during  recent  years  to 
considerable  intensive  study  in  human  medicine,  from 
which  activity  some  interesting  and  useful  information 
has  been  obtained  as  to  its  genesis  and  effect  upon  the 
function  of  the  lung.  This  condition  is  of  course  a  purely 
environmental  one,  the  degree  and  particular  kind  of 
''  dusting  "  being  dependent  upon  the  duration  and 
nature  of  exposure  of  the  particular  individual.  This 
Garden  is  situated  beside  an  active  railroad  trunk  line  so 
that  the  opportunity  for  coal  dust  inhalation  is  con- 
tinuous. The  degree  of  anthracotic  pigmentation  of  the 
lungs  and  related  serous  membranes  is  really  negligible 
and  with  one  questionable  exception,  we  have  not  seen 
fibrosis  due  to  this  cause  in  any  animal.  The  one  excep- 
tion, an  amazon  presented  and  living  in  the  Garden  but 
three  months,  at  necropsy  showed  an  interstitial  chronic 
bronchitis  and  pneumonitis  stretching  out  from  the  hilum, 
all  of  the  affected  area  being  deeply  pigmented.  The 
picture  was  comparable  to  what  might  be  expected  from 
a  second  degree  anthracotic  fibrosis  of  Landis  and  Pan- 
coast.     Many  specimens  come  to  autopsy  with  some  grade 


160  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

of  pigmentation,  but  none,  except  the  one  above,  vnth. 
resultant  fibrosis.  The  degree  of  anthracosis  is  usually 
so  slight  that  it  has  been  considered  important  enough  to 
include  in  the  diagnosis  but  eighteen  times  and  curiously 
enough  seventeen  of  these  were  observed  in  birds.  Were 
there  more  cases  it  might  be  profitable  to  plot  their  exhi- 
bition period  but  the  use  of  this  small  number  might  lead 
to  error ;  the  average  leng-th  of  exhibition  of  the  birds  was 
about  a  year.  It  is  common  to  observ^e  some  black  speck- 
lings  of  the  air  sacs,  as  if  pepper  were  dusted  on  them  as 
has  been  said  before,  but  even  this  is  rarely  marked.  It  is 
most  often  seen  in  the  Anseres,  Psittaci  and  Struthiones 
but  a  goodly  number  of  cases  occur  in  the  long- 
lived  Passeres.  ■ 

The  distribution  of  the  pigment  is  essentially  the  same 
throughout  Mammalia — peribronchial,  submucous  and  in 
the  Ij^mph  nodes  at  the  root  of  the  Imig.  In  the  birds  it  is 
first  seen  in  the  subepithelial  spaces  of  the  septa  of  the 
small  alveoli  where  they  project  into  the  secondaries, 
later  accumulating  in  the  connective  tissue  of  the  main 
septa.  Collections  under  the  pleura  and  at  the  root  of 
the  lung  are  rare,  the  dust  usually  spreading  out  along  the 
air  passages  into  the  air  sacs. 

Other  forms  of  pneumonokoniosis  are  unkno^vn. 
Although  animals  must  inspire  much  dust  from  dry  feed 
and  from  floors  it  must  be  caught  early  and  removed  by 
snorting  or  by  the  lymphatic  drainage.  It  seems  fairly 
well  accepted  that  dusts  are  dangerous  to  the  degree  that 
they  contain  inorganic  substance  and  as  these  animals  are 
not  exposed  to  concentrated  mineral  or  metallic  dusts,  no 
effects  are  seen. 

Infarction  of  Lung. 

Infarctions  of  the  lung,  while  not  at  all  common,  are 
interesting  because  of  their  incidence  in  the  Carnivora 
and  in  the  distribution.  The  figures  concern  the  mammals 


THE  RESPIRATORY  SYSTEM  161 

only  since  the  decision  for  or  against  infarct  in  the  birds  is 
very  difficult  because  of  the  frequency  in  this  class  of 
hemorrhage  A\4th  pulmonary  congestion.  There  were 
eleven  single  or  double  non-septic  infarcts,  of  which  seven 
occurred  in  Camivora,  one  in  an  ungulate,  two  in  Pri- 
mates and  one  in  a  rodent.  The  existence  of  parasites  was 
excluded  in  most  of  the  cases  but  could  not  be  entirely  in 
all.  Eight  of  these  infarcts  were  on  the  left  side,  five  of 
these  being  in  the  lower  lobe. 

Emphysema. 

Emphysema  of  the  atrophic  and  chronic  vesicular 
types  with  the  soft,  fluffy,  pigmented  or  pale  pink  organ 
has  not  occurred  in  the  animals  under  observation.  Acute 
vesicular  emphysema,  such  as  is  seen  in  chronic  bronchial 
and  cardiac  diseases,  has  been  encountered  several  times. 
Cardiac  lesions  were  found  four  times,  nephritis  eleven 
times,  acute  enteric  conditions  seven  times,  hepatic  dis- 
eases seven  times.  Two  cases  of  wide  spread  amyloid 
disease  showed  a  deposit  of  this  substance  in  the  alveolar 
walls.  It  is  quite  common  to  find  some  grade  of 
emphysema  in  monkeys  dying  from  osteomalacia  and 
rickets.  The  process  is  then  most  prominent  in  the  upper 
lobes  and  along  the  free  anterior  margins.  The  incidence 
in  the  orders  is  Primates  5,  Lemures  2,  Camivora  2, 
Pinnipedia  (drowning)  1,  Rodentia  1,  Ungulata  4,  Mar- 
supialia  4.  The  best  example  was  found  in  a  Skunk 
{Mephitis  mesomelas)  having  a  general  infection,  ema- 
nating from  the  cranial  sinuses,  and  cardiac  dilatation. 
Emphysema  does  not  seem  to  occur  in  birds  for  only  one 
was  seen  which  seemed  to  present  this  condition.  This 
was  a  Bald  Eagle  {Ealiceetus  leucocephalus)  with  chronic 
renal  and  enteric  disease  and  cardiac  hypertrophy.  The 
lungs  were  tensely  distended  under  their  serous  covering 
and  showed  a  few  small  bull«  anteriorly.  Unfortunately 
a  histological  preparation  is  not  at  hand. 


162   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Tumors. 

Tumors  of  the  lung  are  moderately  common,  both  of 
primary  and  secondary  origin.  Thus  we  have  seen  three 
primary  and  six  metastatic  growths  in  mammals  and  one 
of  each  kind  in  birds.  The  primaries  were :  carcinomata 
in  a  civet  {Viverra  tangalunga),  a  bandicoot  {Thylacomys 
lagotis),  a  kangaroo  {Macropus  rufus),  and  a  lorikeet 
{Glossopsittacus  concimius) .  The  secondaries  were  :  car- 
cinomata in  a  black  bear  {Ursus  americanus),  a  polar  bear 
(Ursus  maritimus),  a  lion  (Felis  leo),  and  a  dasyure 
{Dasyurus  maculatus) ;  sarcomata  in  a  prairie  wolf 
{Canis  latrans),  and  a  raccoon-like  dog  {Canis  pro- 
cyonoides) ;  adenocarcinoma  in  a  chestnut-eared  finch 
{Amadina  castanotis). 

The  histological  character  of  the  primary  cancers 
would  place  them  in  group  of  the  nodular  and  infiltrative 
types  of  Kauffman.  They  all  seem  to  have  taken  their 
origin  from  the  smaller  bronchi,  the  usual  starting  point. 
The  growths  were  small  in  the  civet  and  bandicoot  and 
strongly  suggest  that  the  tumors  arose  in  bronchi  occu- 
pied by  parasites ;  such  bodies  could  not  be  demonstrated. 
It  is  the  usual  tiling  to  find  in  cases  of  parasitism  of  the 
bronchi  that  if  there  be  no  ulcerative  destruction  of  tissue 
the  epithelium  undergoes  some  form  of  hyperplasia,  and 
even  structural  metaplasia  in,  the  air  tubes  supplied  with 
cuboidal  or  cylindrical  cells.  Epithelia  many  layers  deep 
have  been  encountered,  usually  arranged  in  orderly 
fashion  but  frequently  ''  papillomatoid, "  suggesting  the 
epidermal  layers  yet  not  so  far  as  to  show  protoplasmic 
bridges.  Distention  of  various  degrees,  affected  by  the 
contents  of  the  tube  and  the  surrounding  inflammation, 
are  common.  Such  a  picture  naturally  resembles  epi- 
thelioma and  indeed  growths  of  this  nature  are  reported 
as  due  to  verminous  pneumonitis. 

There  are,  especially  in  cats  and  dogs,  small  scattered 
adenomatoid  groAvths(2)   under  the  pleura  and  in  the 

(2)    Ball,  Jour.  Vet.,  1907. 


THE  RESPIRATORY  SYSTEM  163 

pulmonary  substance,  thought  to  originate  in  the  alveolar 
epithelium  and  occasionally  growing  to  large  size;  the 
case  in  the  kangaroo  may  have  had  this  origin.  It  ivas 
the  only  primary  tumor  to  give  metastasis  (to  the  spleen 
and  gastric  wall),  the  secondaries  being  decidedly  ade- 
nomatous in  character. 

Metastatic  groAvths  come  from  the  following  originals : 
two  from  the  thyroid,  well  known  to  give  pulmonary 
embolism  in  dogs;  one  each  from  the  breast,  uterus, 
adrenal,  intestine  and  kidney.  The  form  assumed  is  a 
gray  and  red  mass  lying  under  the  pleura  or  an  isolated 
nodule  in  the  substance.  Sarcomatosis,  the  form  appar- 
ently spreading  out  from  the  hilum  and  growdng  in 
isolated  grayish  tubercular  masses,  has  not  been  seen. 

The  Pleuea. 

The  pleura  is  a  tissue  apparently  quite  susceptible  to 
infection  in  mammals  and  so  closely  associated  with  the 
air  sacs  in  birds  as  to  be  a  part  of  the  same  membrane, 
therefore  the  two  being  affected  together.  Throughout 
the  higher  class  all  orders  give  copious  examples  of  the 
involvement  of  the  pleura,  principally  of  course  as  an 
accompaniment  or  a  sequel  to  pneumonitic  or  bronchitic 
processes  but  also  as  a  part  of  acute  infectious  diseases, 
such  as  hemorrhagic  septicemia,  pleuropneumonia  and 
the  like.  However  two  orders  present  such  a  number  of 
instances  of  pleuritis  that  they  deserve  notice.  The  seals, 
Pinnipedia,  of  w^hicli  we  have  twenty  autopsy  records, 
showed  inflammation  of  this  membrane  four  times,  three 
of  which  were  dependent  upon  pulmonary  infection  and 
one  apparently  due  to  general  septicemia  with  trifling 
damage  to  the  lung  proper.  One  of  the  first  cases  had 
gone  on  to  empyema  of  the  classical  type,  a  shrivelled  dry 
almost  carnified  lung  with  a  thick  fibrinopurulent  cover- 
ing. The  lung  of  the  seal  is  well  divided  into  lobules,  the 
external  surface  being  generously  supplied  with  lym- 
phatic channels  under  the  pleura,  an  arrangement  which 


164  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

should  carry  away  infection  one  would  think.  Perhaps 
this  high  percentage  of  pleurisy  in  our  Pinnipedia  is  but 
accidental.  The  marsupials,  while  having  a  notable  per- 
centage of  pleurisy  both  among  all  the  cases  and  in  rela- 
tion to  the  number  of  postmortems,  are  not  so  striking 
from  the  etiological  standpoint  since  practically  all  of 
these  have  suffered  with  Kangaroo  mycosis  or  pneumonia. 
In  over  half  the  cases  of  tliis  infectious  disease  some  grade 
of  pleuritic  exudate  has  been  observed,  only  one,  however, 
going  to  the  stage  of  empyema. 

;  One  cannot  speak  so  definitely  of  pleuritis  in  birds 
since  this  tissue  merely  represents  in  them  the  covering 
of  the  lung  and  is  firmly  attached  posteriorly  to  the  ribs 
and  anteriorly  to  the  air  sacs.  Exudates  show  as  col- 
lections upon  the  air  sac  side  of  the  combined  membrane, 
pleuritis  proper  in  birds  being  an  infiltrative  affair  com- 
ing through  the  pulmonary  tissue  and  therefore  being  a 
part  of  pneumonitis.;  I  notice  a  tendency  in  a  few  articles 
to  write  of  pleuritis  when  the  process  is  confined  to  the 
thorax  but  this  gives  the  impression  that  the  disease  is 
peculiar.  There  seems  no  difference  in  the  gross  and 
minute  appearance  between  thoracic  serositis  and  pan- 
serositis. The  course  of  procedure  seems  to  be  from  the 
anterior  or  mesial  pulmonary  ostia  into  respectively  the 
cervical  and  thoracic  air  sacs  and  this  seems  to  hold  good 
whether  the  infection  be  mycosis  or  fowl  cholera  or  fowl 
pest.  There  are  records  of  104  cases  of  serositis  in  birds 
of  which  45  were  among  parrots,  the  remainder  being 
well  distributed  among  the  various  orders ;  only  two  each 
occurred  in  Galli  and  Anseres,  orders  prominently 
affected  under  domestication.  This  high  percentage  of 
pleuroperitonitis  among  parrots  and  their  congeners  can 
only  be  explained  upon  the  ground  of  a  continued  infec- 
tion of  our  stock  by  the  virus  of  fowl  cholera  and  by 
mould.  One  case  of  undoubted  fowl  cholera  occurred 
recently  and  as  the  records  are  reviewed  a  few  are  dis- 
covered where  the  organism  was  found.     The  virus  must 


HA  t. 


•<Hi 


Fig.   12.— endothelioma  OF  PLEURA.     LEOPARD   (FELIS  N'EBULOSA). 


Fio.   1?. — ENDOTHELIOMA  OF  PLEURA   IN   FIG.    12.      DETAIL  OF  ONE  OF  THE  WARTY   EXCRESCENCES. 


THE  RESPIRATORY  SYSTEM  165 

be  of  low  grade  for  we  have  had  no  severe  and  devasting 
epizootic.  Mycosis  is  constantly  with  us  no  matter  what 
we  do  in  hygienic  measures.  There  was  a  small  group 
of  cases  of  pulmonary  and  serous  membrane  mycosis  com- 
bined mth  staphylococcus  infection  which  carried  off  six 
birds.  The  pathology  of  tliis  group  was  interesting 
because  one  could  follow  the  infection  of  the  bacterium. 
The  anterior  pulmonary  ostium  was  surrounded  or 
covered  by  a  mycotic  mass  and  spreading  downward 
from  this  was  a  grayish  yellow  turbidity  of  the  air  sac 
walls  with  a  delicate  sticky  or  almost  mucilaginous 
exudate  extending  into  the  lateral  abdominal  and  pos- 
terior sacs. 

There  is  on  record  one  tumor  of  the  pleura,  an  endo- 
thelioma, in  a  Clouded  Leopard  (Felis  nehulosa).  It  was 
the  usual  plate-like  thickening  with  warty  excrescences. 
No  metastases  occurred.  (Figs.  12  and  13). 


SECTION  VII 

THE  ALIMENTARY  TRACT.    PART  1.— PHARYNX, 
ESOPHAGUS,  STOMACH  AND  INTESTINES 

The  portion  of  the  anatomy  that  we  now  approach 
varies  in  construction  almost  as  much  through  the  animal 
kingdom  as  do  the  external  shape  and  covering  of  the 
various  orders  and  much  more  than  do  the  other  systems. 
The  reason  for  this  is  obvious,  an  arrangement  accommo- 
dative to  the  differing  food  chiefly,  but  not  a  little  to  the 
ease  with  which  animals  obtain  and  assimilate  their  nutri- 
ment. It  would  be  impracticable  to  describe  all  the  varia- 
tions of  the  orders  discussed  in  this  study,  but  since 
certain  gross  and  minute  differences  are  of  importance  in 
comparative  pathology  they  mil  be  discussed  at  the  ap- 
propriate places.  It  is  my  purpose  to  present  in  a 
subsequent  section  a  discussion  of  food  in  captivity  from 
the  standpoint  of  its  quality  and  quantity  in  relation 
to  pathology. 

(^Doubtless  the  quality  of  food  is  the  largest  factor  in 
the  production  of  disease  both  of  the  alimentary  tract  and 
elsewhere,  but  I  am  not  at  all  sure  that  the  quantity  may 
not  be  equally  important,  in  certain  groups  at  least.  Thus, 
for  example,  the  ungulate  has  nearly  always  available  in 
bedding  a  substance  that  he  can  and  will  eat,  and  the 
prevalent  idea  that  an  animal  will  eat  only  as  much  as  is 
good  for  him  seems  not  to  hold  at  all  times,  since  over- 
filled rumens  are  only  too  common.  It  might  be  thought, 
however,  that  capti\4ty  creates  a  sort  of  pica,  or  that 
enforced  idleness  is  conducive  to  gorging.  The  use  a  few 
lines  above  of  the  word  "doubtless"  may  have  arrested 
the  attention  of  some,  yet  when  the  whole  subject  is 
reviewed  it  seems  entirely  justified.  Plimmer  puts  incor- 
rect food  at  the  head  of  the  list  of  the  causes  of  enteritis, 

166 


THE  ALIMENTARY  TRACT  167 

Brooks  emphasizes  the  importance  of  certain  grasses  and 
musty  fodder,  systematic  writers  detail  among  the  princi- 
pal causes  of  gastritis  spoiled  food,  and  in  zoological 
gardens  specific  disease  like  hog  cholera  and  entero- 
hepatitis  are  relatively  uncommon  while  non-specific 
gastroenteritis  is  the  most  frequent  diagnosis  in  causes 
of  death. 

The  other  factors  to  which  enteritis  is  ascribed  are 
animal  and  vegetable  parasites  and  mechanically  oper- 
ative foreign  bodies,  the  last  being  unimportant.  Just 
how  important  the  first  mentioned  are  is  a  matter  of  some 
question  which  must  be  subjected  to  considerable  study 
before  any  solution  can  be  expected. 

If  for  no  other  reason  than  that  the  gross  and  minute 
pathological  anatomy  of  gastro-entero-colitis  is  the  same 
through  the  mammals  and  birds  (aside  from  a  few 
specific  lesions  like  enterohepatitis,  typhoid  fever,  etc.), 
while  the  food  and  bacteria  var^^  it  would  seem  probable 
that  the  ultimate  cause  is  the  same,  a  poison  which  can  be 
formed  alike  in  the  carnivorous  and  herbivorous  gut,  and 
not  dependent  upon  bacteria,  but  upon  the  chemistry  of 
the  food  or  of  the  intestinal  mucosa.  To  put  the  matter 
more  simply,  the  lesions  being  the  same  under  nearly  all 
conditions  is  not  the  cause  the  same,  and  is  it  not  a 
poisonous  product  from  food  or  the  intestinal  lining.  It 
is  profitable  here  only  to  mention  the  marked  similarity 
of  enteric  lesions  under  the  differing  conditions  and  in 
different  orders.  We  shall  study  chartwise,  the  various 
forms  of  inflammation  from  the  cardia  to  the  anus  in 
terms  of  their  anatomical  diagnosis  and  most  probable 
etiology  in  an  attempt  to  throw  light  upon  the  matter,  and 
later  present  the  physiology.  System  requires,  however, 
some  attention  first  to  anatomical  order  so  that  a  brief 
review  of  the  esophageal  and  pharyngeal  conditions 
is  indicated. 


168  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Pharynx. 

The  buccal-pharyngeal  cavity  in  mammals  is  used 
chiefly  as  a  passage  way  for  food  and  as  the  place  where 
some  of  them  triturate  and  insalivate  the  bolus.  Certain 
orders,  Primates,  rodents  and  marsupials,  use  this  cavity 
thoroughly  at  the  first  mastication,  others  use  it  little  at 
first  but  may  ruminate,  the  ungulates,  while  strict  car- 
nivores use  it  very  little.  ^  The  Aves  use  their  pharynx 
almost  exclusively  as  a  passage,  and,  despite  the  presence 
of  a  certain  amount  of  salivary  gland  tissue,  probably  do 
not  digest  any  substance  in  this  cavity.  The  crop  or 
ingluvies  is  a  sac  to  permit  of  salivary  digestion  but  is 
really  a  storehouse  to  allow  rapid  feeding  without  over- 
filling of  the  proventricle.  The  esophagus  extends  from 
the  pharynx  to  the  cardiac  opening  of  the  stomach  in 
mammals  and  widens  into  the  glandular  stomach  or 
proventricle  in  birds,  the  upper  end  of  wliich  lies  in  front 
of  the  lower  third  of  the  left  lung  behind  the  heart. 

Inflammations  of  the  buccal,  pharyngeal  and  esopha- 
geal walls  are  relatively  common  in  certain  orders  espe- 
cially ground  birds  and  grazing  ungulates.  This  would 
seem  to  be  explained  on  the  basis  of  injury  to  the  mucosa 
by  sharp  or  pointed  objects  picked  up  while  feeding.  The 
character  is  usually  necrotizing,  but  need  not  be,  and  the 
bacteriology  is  not  specific.  Definite  infectious  diseases 
like  diphtheria  and  actinomycosis  are  not  included  here, 
but  it  might  be  mentioned  that  the  second  disease  cited 
is  believed  to  be  started  by  the  penetration  of  the  organ- 
isms into  wounds  made  by  sharp  grasses.  Certain  orders, 
notably  Ungulata,  Passeres,  Psittaci  and  Struthiones,  are 
quite  susceptible  to  mycotic  infestation  and  we  have  seen 
an  outbreak  of  thrush  in  Kites  (Accipitres).  It  is,  how- 
ever, interesting  and  possibly  significant  of  peculiar 
protective  powers  in  the  upper  alimentary  tract,  that 
strict  carnivores  have  failed  to  show  ulcerative,  purulent 
or  necrotizing  inflammations  of  the  mucosa  from  the 


THE  ALIMENTARY  TRACT  169 

mouth  to  the  cardia.  There  has  been  no  important  data 
upon  ingluveal  indigestion  or  esophageal  obstruction. 
(Birds  especially,  and  occasionally  mammals,  gorge  them- 
selves or  take  too  large  a  bolus,  but  it  seems  as  if  this  is 
only  fatal  where  some  distinct  important  pathology  is 
present  which  has  reduced  their  resistance.  In  the  lower 
esophagus  one  has  to  deal  with  worms  in  connection  with 
the  proventricle  in  birds,  but  no  mammals  seem  to  have 
suffered  with  temporary  or  permanent  strictures.  *'  Crop- 
binding  ' '  has  occurred  in  the  following  orders :  Psittaci, 
GalH;  and  overfilling  of  the  esophageal  dilatation  was 
seen  in  Accipitres  and  Herodiones ;  Columbidse  with  their 
double  crop  were  not  affected  by  this  abnormal  collection 
of  food  in  the  esophagus. 

Dilatations  of  Esophagus. 

The  mammals  have  shown  three  dilatations  of  the 
esophagus  interesting  enough  to  detail  briefly : 

Mongoose  Lemur  (Lemur  mongoz)  6  .  Sacculo-fusiform  dilatation 
of  esophagus,  probably  congenital,  with  adjacent  fibrosis  of  lung.  In 
poor  condition  for  several  years  but  recovered  satisfactorily  from  a 
bad  cut  inflicted  by  cage-mates.  At  autopsy  the  general  condition  is 
poor,  hair  missing  in  spots,  all  skin  dry  and  atrophic  with  patches  of 
keratotic  dermatitis.  All  tissues  anemic,  muscles  lusterless.  Right 
lung  collapsed,  brown  and  pink,  spotted  with  anthracosis.  Left  lung 
pushed  forward  and  to  left  by  a  mass  in  the  posterior  mediastinum. 
Lower  lobe  in  its  posterior  portion  is  adherent  to  esophageal  mass. 
Lower  half  of  this  lobe  beginning  where  bronchus  ends  and  extending 
over  anterior-posterior  surfaces  shows  marked  fibrotic  processes  and 
at  one  point  in  tissue  between  end  of  bronchus  and  adherent  esophagus 
there  is  no  lung  tissue  remaining.  No  recent  consolidations.  Bronchial 
lymph  nodes,  small,  firm  homogeneous  pale  brown  with  specks  of  anthra- 
cosis. Heart  conti'acted,  normal  in  size,  firm  red-brown.  Aorta  is 
firmly  adherent  to  esophageal  mass  where  bronchus  crosses  it.  The 
lower  half  of  the  esophageal  from  the  hilum  of  the  lung  to  the  cardia 
is  the  seat  of  a  dilatation,  fusiform  for  the  most  part,  but  with  a  saccular 
portion  anteriorly.  This  latter  presses  the  left  bronchus  upward  and 
heart  forward.  The  wall  of  the  tube  is  slightly  irregularly  thickened 
but  there  is  no  cicatrix  and  mucosa  shows  slight  hypertrophic  condi- 
tion. A  large  mass  of  food  occupies  the  dilatation.  Stomach  is  empty 
save  for  gas.  Mucosa  is  soft,  smooth,  pale  pink.  Duodenum  shows 
12 


170  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

slight  swelling  of  the  rather  pale  yellow  submucosa  and  mucosa,  but 
the  tips  of  the  villi  are  injected.  Intestine  contains  only  a  little  slimy 
mucus.  Large  intestine  contains  a  mass  of  constipated  feces.  The 
esophageal  dilatation  seems  to  have  been  congenital  although  it  is  barely 
possible  that  the  fibrosing  pneumonitis  and  pleuritis  may  have  aided 
and  caused  it  by  traction.  It  has  been  doubtless  the  cause  of  the  ani- 
mal's inanition. 

Black  Bear  6  {Ursus  americanus).  Sacculate  dilatation  of  esophagus 
with  chronic  esophagitis.  Chronic  hypertrophic  gastritis.  Chronic  lym- 
phadenitis. Fatty  degeneration  of  liver.  Acute  catarrhal  enteritis.  Had 
been  vomiting  more  or  less,  nearly  every  day  for  two  months  and 
did  not  eat  for  six  days  before  death.  Mouth,  pharynx  and  esophagus 
are  full  of  macerated,  unrecognizable  food.  Pharnyx  seems  normal. 
Larynx  is  yellow,  mucosa  rough  and  slightly  thickened  in  places  espe- 
cially just  above  the  vocal  chords.  No  excess  of  mucus.  Esophagus 
in  neck  is  dilated.  Mucosa  is  rough,  irregular  yellow-brown.  This 
dilatation  proceeds  downward  so  that  at  thoracic  opening  tube  is  twice 
normal  size.  Upon  entering  thorax  this  dilatation  turns  to  right  and 
in  in*egular  saccular  form  extends  to  diaphragm  compressing  heart  and 
lungs  to  left.  The  main  course  of  it  then  recrosses  the  midline,  aorta 
being  slightly  twisted  as  it  regains  position  in  front  of  vertebrae.  The 
wall  is  irregularly  thickened  from  pseudomembranous  patches  and 
some  hypertrophy  of  mucous  and  submucous  layers.  Muscle  and  serous 
coats  are  considerably  thinned.  The  sac  is  full  of  macerated  food  and 
gas.  The  right  lung  is  compressed  small  resilient  red-gray.  No  ad- 
hesions in  either  pleura.  On  section  the  lung  tissue  is  found  to  be 
slightly  edematous,  red  gray,  compact  and  while  not  atelectatic  yet 
crepitates  much  less  than  noi-mal.  Left  lung  is  the  seat  of  passive  con- 
gestion in  lower  lobe  and  lower  half  of  upper  lobe.  The  upper  half  of 
lobe  is  compressed  and  subcrepitant  like  the  right  lung.  Lymph  glands  of 
neck  and  thorax  are  much  enlarged  firm  with  large  irregular,  clear  out- 
lined follicles  and  brown  firm  homogeneous  pulp.  The  aorta  shows 
slight  roughening,  the  intima  being  smooth  and  homogeneous. 

Lion  6  {Felis  leo).  Ten  and  one-half  months  old.  Cystic  paren- 
chymatous goitre,  dilatation  of  esophagus,  ulcerative  enteritis.  Acute 
glomerular  nephritis.  Chronic  hyperplasia  of  spleen.  Fatty  infiltration 
of  liver.  Bronchopneumonia  (from  pressure  of  goitre).  Ascaris  in  in- 
testines. Had  lump  on  neck  for  several  weeks,  ate  very  little  and 
seemed  to  have  hard  work  to  get  anything  down.  Stopped  eating  to- 
ward the  last  and  vomited  water  and  foam.  There  is  a  small  ulcer  with 
everted  lips  just  below  left  incisor  on  loAver  lip  which  the  keeper  says 
is  of  several  months'  duration.  The  thyroid  is  much  enlarged  and  forms 
a  large  mass  in  the  upper  chest  and  extends  far  up  in  the  neck.  Because 
of  this  mass  the  lungs  are  pushed  far  down  in  the  chest.  Heart  also 
lies  very  low.  The  thyroids  are  enormously  enlarged  and  cystic,  the 
right  measuring  13  x  8.5  x  5  cm.  and  the  left  19  x  9  x  5  cm.     The  lungs 


Fig.  14. — DILATATION  OF  E^OF11AOL^.  LION  (FKLKS  LEO).  DII.ATAIION  PROBABLY  DUE 
TO  OBSTRUCTION  BY  ENLARGED  THYROID  BODY.  IN  ILLLSTR.AFION  DLSFENFION  OF  ESOPHA- 
GUS CAN    BE  SEEN    IN  THE  FORK  OF  THE  THYROID  LOBES. 


THE  ALIMENTARY  TRACT  171 

are  pale  pinkish  white.  Air  content  increased  in  places,  decreased  in 
others.  No  hypostatic  congestion.  The  lungs  seem  normal  except  at 
the  apex  where  they  are  collapsed  probably  from  pressure  on  lung  by 
enlarged  thyroid  which  dips  down  into  the  chest  for  at  least  three 
inches  filling  entirely  the  apex  of  the  chest.  One  bronchial  lymph  gland 
was  about  the  size  of  a  walnut,  the  rest  were  normal.  The  heart  seems 
normal  except  for  its  slightly  low  position.  The  abdomen  contains  about 
300  c.c.  of  deep  yellow  highly  albuminous  fluid.  No  adhesions.  The 
liver  is  softer  than  normal,  glistening,  smooth,  moist  and  very  yellow 
particularly  at  the  edges.  The  gall-bladder  contains  a  green  mucoid 
bile  and  the  duct  is  patulous.  Spleen  and  kidneys  are  normal.  Mouth 
and  pharynx  are  normal.  The  esophagus  is  much  dilated  above  the 
thyroid.  The  enlarged  thyroid  pressing  upon  it,  has  acted  as  a  distinct 
obstruction.  In  this  pouch  was  a  large  amount  of  food  probably  (from 
the  histoiy)  eaten  two  days  before.  The  esophagus  below  this  point 
was  normal  except  for  the  presence  of  thin  mucus.  Stomach  empty 
save  for  two  small  bits  of  meat.  The  duodenal  walls  are  much  thickened, 
mucosa  covered  with  small  ulcers  many  with  a  hemorrhagic  base;  there 
were  present  also  a  feAV  small,  round  worms. 

Here  are  presented  three  different  dilatations,  the 
first  probably  congenitally  started  and  aided  by  pul- 
monary fibrosis,  therefore  secondarily  a  traction 
diverticulum,  the  second  probably  entirely  congenital, 
and  the  unusual  third  case  due  to  obstruction  by  an 
enlarged  thyroid.  'In  this  connection  might  be  mentioned 
small  saccular  diverticula  in  the  proventricle  of  a  Fire 
Finch  {Lagonosticta  senegala),  and  at  the  pylorus  in  a 
Puma  {Felis  concolor).  Neither  of  these  seemed  of  any 
significance  and  played  no  part  in  the  death  of  the 
animals ;  they  did  not  seem  to  be  artificially  produced,  by 
worms,  for  example. 

The  Proventricle. 

'v  The  proventricle  or  forestomach  of  birds,  is  the  seat 
of  active  secretion  of  the  gastric  juice  in  nearly  all  orders, 
although  Jobert  believes  that  the  mucosa  of  the  gizzard 
may  contribute  some  digestive  fluid,  and  there  are  active 
glands  in  this  tissue  in  a  few  orders.  The  proventricle 
does  not  act  as  a  reservoir  during  digestion,  but  as  soon 
as  the  juices  are  well  mixed  mth  the  bolus  the  food  is 


172   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

passed  on  to  the  gizzard.  The  organ  has  a  rather  free 
position,  at  least  as  far  as  its  left  lateral  and  downward 
movements  are  concerned  for  the  left  lateral  abdominal 
air  sac  is  free  on  that  side  of  the  gastric  complex  and  the 
left  lateral  thoracic  also  extends  do-\vn  the  side  of  the 
proventricle.  Upward  and  anteriorly  is  the  heart.  Some 
of  the  cases  of  proventricular  spiropteriasis  have  showTi 
very  marked  congestion  of  the  left  lung,  possibly  due  to 
the  closure  of  the  anteroinferior  air  sac  aperture  on  that 
side.  Aside  from  parasitism,  affections  of  this  organ  are 
not  very  numerous?) 

Infestation  with  spiroptera  and  with  less  dangerous 
worms  was  quite  serious  at  one  time,  but  since  routine 
examinations  of  all  suspicious  birds  has  been  practiced 
this  parasitism  has  been  under  control.  This  is  a  subject 
of  especial  importance  to  collections,  and  will  be  described 
in  a  special  section.  The  susceptibility  of  the  proventricle 
to  damage  seems  from  our  figures  to  be  very  distinctly  a 
matter  of  zoological  order.  In  so  far  as  parasites  are 
concerned,  the  parrot  group  stands  away  ahead  of  all 
others  having  an  incidence  among  autopsies  upon  Psittaci 
of  16  per  cent. ;  after  them  come  the  Picariae  with  9  per 
cent.,  and  Columbse  and  Passeres  each  with  4  per  cent. 
AVhen,  however,  non-verminous  conditions  are  reviewed 
the  anserine  birds  are  found  the  most  susceptible,  3  per 
cent,  of  the  autopsies  upon  this  order  revealing  proven- 
tricular lesions;  after  them  come  the  Columbae,  2  per 
cent.,  and  Psittaci,  1.5  per  cent.  In  this  group  are 
included  inflammations  of  all  kinds,  dilatations  and  dis- 
tentions and  some  lesser  matters. 

\  This  part  of  the  stomach  has  been  involved  in  our  cases 
of  fowl  diphtheria,  showing  a  distinct  mucopurulent 
inflammation  with  penetration  into  the  depths  of  the 
glands ;  no  separation  of  the  mucosa  occurred,  but  ulcera- 
tion was  seen.  Perforation  was  observed  thrice,  tmce  by 
ulceration  around  a  worm  while  it  was  boring  into  the 


THE  ALIMENTARY  TRACT  173 

muscular  layer  and  once,  in  an  ostrich,  by  the  penetration 
of  a  nail.  Obstruction  of  this  division  of  the  stomach  by 
impaction  of  food  and  stones  is  not  common,  but  does 
occur.  The  reason  for  such  obstruction  is  usually  very 
obscure.  Sometimes  it  seems  due  to  the  feeding  of 
seeds  and  the  like  in  too  finely  divided  form,  whereby  food 
and  pebbles  are  taken  up  together.  Some  of  the  smaller 
birds  have  had  in  times  past  too  many  small  pebbles  in  the 
cages,  while  others  have  had  too  large  seeds,  thus  appar- 
ently trying  to  use  the  pebbles  to  crush  them.  It  would 
seem  also  that  the  birds  had  really  eaten  too  much  and 
could  not  accommodate  it  in  the  gullet  and  gizzard ;  this 
seems  surely  true  in  three  or  four  Accipitres.  Most  often, 
however,  we  have  had  to  fall  back  upon  the  inadequate 
explanation  of  pica  or  perverted  appetite. 

i  Impacted  proventricles  and  gizzards  have  been 
observed  thirty-four  times,  in  eight  of  which  it  seemed 
the  sole  cause  of  death,  and  therefore  probably  entirely 
due  to  foreign  bodies  in  food.  The  theory  is  accepted 
that  dilatation  and  obstruction  will  not  occur  if  the  motor 
power  of  the  gastric  wall  be  normal  and  no  inflammation 
exist.  In  this  regard  we  can  only  discover  five  birds  (the 
mammals  will  be  discussed  later)  ^vith  any  distinct  inflam- 
matory or  degenerative  disease  of  this  part  of  the 
anatomy  and  two  with  lesions  elsewhere  which  might 
affect  the  musculature;  this  leaves  the  vast  majority  of 
gastric  obstruction  in  birds  unexplained  on  basis  of 
defective  motor  power,  therefore  probably  dependent 
upon  the  character  of  material  consumed.  The  anserine 
birds  and  parrots  are  most  often  affected  by  this  form 
of  obstruction. 

i,  Acute  or  chronic  dilatation  of  the  forestomach  and 
gizzard  is  very  rare  in  birds,  it  ha\ing  occurred  only 
thrice  in  our  records,  a  finch,  a  parrot  and  an  owl;  the 
causes  were  entirely  unknown  since  the  cavities  were  not 
overfilled  Avith  food.  / 


174  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

The  Stomach. 

Impaction  of  the  mammalian  stomach  is  a  diagnosis 
made  but  once  in  our  records,  an  Indian  Antelope  {Anti- 
lope  cervicapra) ,  and  this  is  viewed  with  suspicion.  The 
rumen  was  undoubtedly  tightly  packed,  being  distended  to 
its  fullest  capacity  with  rather  dry  and  not  properly 
softened  grain.  However,  postmortem  changes  had 
advanced  and  therefore  observations  in  the  whole  body 
were  not  dependable.  The  bulk  of  food  w^hich  can  be 
accommodated  by  the  rumen  is  very  large,  and  had  this 
not  been  dry  and  firm  the  condition  might  not  have  been 
interpreted  so  seriously.  Many  animals  come  to  autopsy 
with  a  well  filled,  indeed  even  mth  a  well  packed  stomach, 
but  there  is  usually  sufficient  reason  for  this  or  there  is 
distinct  pathology  to  account  for  death. 

All  this  of  course  impUes  a  stomach  of  normal  or 
approximately  normal  dimensions  since  distention 
beyond  this,  or  dilatation  of  the  stomach,  is  more  definite. 
In  veterinary  medicine,  gastric  tjanpanites  (rumen  alone 
or  all  stomachs,  or  the  simple  stomach)  is  ascribed  to 
food  that  ferments  easily  when  taken  in  excess  or  in  the 
presence  of  defective  motor  power,  to  constrictions  by 
scars  and  to  obstructive  tumors;  excessive  feeding  is 
sometimes  mentioned  but  given  a  subordinate  role.  Our 
records  throw  very  little  light  upon  the  subject  since  only 
five  cases  were  observed.  Four  of  these  five  seem  to  be 
due  to  acute  fermentation  independent  of  gross  physical 
obstruction,  while  one,  a  Cape  Hunting  Dog  {Lycaon 
pictus)  showed  an  old  chronic  ulcerative  gastritis  with 
both  healed  and  active  ulcers  distorting  the  pyloric  end  of 
the  organ.  Three  of  the  first  mentioned  four  were 
monkeys  and  one  was  an  ocelot.  The  stomach  of  the 
voracious  monkey  is  at  time  of  dissection  usually  well 
filled,  but  in  these  cases  there  seems  no  doubt  that  gas  and 
excess  fluid  had  distended  the  cavity  enormously,  in  one 
instance  apparently  assisting  in  acute  cardiac  dilatation. 


Fli 
NTS). 
THE   CARD 


xuVpT'^,"'^'^.^.-.^.?-^'^'^*   '^'  '-^TO-^'ACH   WALL.      CO.\LMO\     OPO 
THLRh     lb      GtNtRAL    MODERATE     CHRONIC      GASTR 


IM    (DIDELPHYS  VIRGIMA- 
WITH      ROLND    ULCERS    NEAR 


THE  ALIMENTARY  TRACT  175 

There  were  no  obvious  reasons  for  assuming  any  damage 
to  the  gastric  motor  mechanism. 

Gastkic  Ulcers. 

Gastric  ulcer,  so-called  peptic  or  round  ulcer  of  the 
stomach,  having  a  chronic  course  and  leading  to  radiating 
scars  of  the  mucosa  is  not  common  in  the  lower  animals, 
but  frequent  enough  in  the  human  being.  The  form  of 
ulcer  in  question  has  at  present  no  adequate  explanation, 
or  at  least  there  is  no  one  cause  which  will  answer  for  all 
cases.  Local  injuries  from  within  or  without  the  stomach, 
bacterial  embolism,  entrance  of  bile  through  the  open 
pylorus  and  many  other  factors  have  been  named  in  the 
causation  but  can  seldom  be  used  in  any  given  case.  In  the 
lower  animals  with  their  relatively  frequent  parasitic 
infestation,  another  factor  is  added.  In  analysis  of  our 
statistics  I  have  separated  ulcerative  gastritis  from  para- 
sitic and  mycotic  ulcerations  and  from  peptic  ulcers ;  the 
first  is  discussed  in  later  paragraphs.  Parasitic  ulcers  of 
the  stomach  occur  chiefly  in  our  native  marsupial,  the 
opossum,  and  in  some  Carnivora ;  physaloptera,  strongy- 
lus,  ascaris,  and  gastrophilus  have  been  found.  The 
kangaroos  are  frequently  affected  (8  cases)  with  an 
acute  or  subacute  ulceration  of  the  gastric  wall,  mthout 
much  general  gastritis.  The  lesion  is  peculiar  in  appear- 
ance. The  youngest  ulcers  are  black  or  dark  gray,  flat 
necroses  of  the  mucosa  alone  and  indeed  the  process  very 
frequently  penetrates  no  deeper.  Older  lesions  spread 
laterally  and  may  be  preceded  by  a  very  narrow  con- 
gested line  but  there  is  no  raised  edge  nor  does  there  seem 
to  be  submucous  infiltration.  If  the  process  be  rapid 
a  loose  dirty  slough  may  form.  Certain  of  the  advanced 
cases  of  Kangaroo  mycosis  mil  present  more  infiltrative 
lesions  of  the  gastric  wall  leading  to  large  and  well 
defined  necrotic  areas ;  they  may  at  times  penetrate  the 
whole  wall  outward.    (See  page  580.) 


176  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

True  peptic  ulcers  have  been  found  in  Primates,  4; 
Carnivora,  5;  Pinnipedia,  2;  Insectivora,  Ungulata  and 
Hyraces  each  one.  The  London  Garden  reports  that  gas- 
tric ulcerations  occur  most  often  in  Carnivora  and 
Marsupialia.  Those  in  the  last  three  orders  of  our  hst 
were  small,  usually  multiple  and  relatively  superficial. 
The  ulcers  found  in  Primates  and  Carnivora  present  the 
usual  pictures  seen  in  man.  In  one  example  in  each  of 
these  orders  radiating  scars  of  healed  defects  are  men- 
tioned in  the  notes.  None  of  them  seems  to  have  led  to 
cancer,  and  in  only  one,  a  wolf  (Canis  lupus),  was  the  scar 
tissue  sufficient  to  cause  definite  impediment  to  the  motil- 
ity of  the  stomach.  Six  of  the  fourteen  examples 
appeared  on  the  greater  curvature,  the  remainder  on  the 
lesser.  Ten  ulcers  were  in  the  pyloric  division,  the  other 
four  being  scattered.  No  other  pathology  is  found  com- 
mon to  these  cases  which  might  be  drawn  into  etiologi- 
cal association. 

Tumors. 

Tumors  of  the  gastric  complex  are  not  at  all  common, 
there  being  only  the  following  to  report:  Primates, 
Hamadryas  Baboon  (Papio  Jiamadryas) ,  diffuse  ade- 
noma; (none  in  Carnivora  with  the  most  ulcers) ;  Mar- 
supialia, Eed  Kangaroo  {Macropus  rufus),  maUgnant 
papilloma  with  metastases.  The  former  tumor,  shown  in 
Fig.  17,  was  a  diffuse  soft  excrescence  beginning  near  the 
pylorus  and  stretcliing  along  the  lesser  curvature  toward 
the  cardia.  Histologically  it  was  made  up  of  glandular 
acini  growing  in  all  directions  but  always  maintaining 
normal  relations  of  cells  and  basement  membrane.  There 
were  no  metastases  and  other  reasons  for  death  existed. 
The  tumor  of  the  kangaroo  stomach  was  a  true  epithelio- 
matous  cancer  with  metastases  to  liver,  spleen,  and  kid- 
ney. Only  one  secondary  tumor  was  observed,  from  a 
carcinoma  of  the  lung  in  a  Red  Kangaroo  (Macro- 
pus  rufus). 


Fit..   16.— MULTIPLE  GASTRIC  ULCERS.     COMMON  WOLF  (CANIS  LUPUS).     CHRONIC  GASTRITIS 
WITH  NUMEROUS  IRREGULAR  ULCERS  OF  THE  PEPTIC  TYPE. 


Vu..    17.  — PAIMM.OMA   Ol-    VroMAeH.      HAMADRVAS    BAHOON    (I'AI'IO    HAMADRYAS). 


THE  ALIMENTARY  TRACT  177 

The  Intestine. 

Inflammation  of  the  gastrointestinocolic  tube  is  the 
most  important  single  condition  with  which  handlers  of 
animals  have  to  deal,  and  unfortunately  it  can  seldom 
be  diagnosed  clinically,  early  and  accurately  enough,  to 
make  treatment  useful.  At  this  Garden  some  evidence  of 
acute  or  chronic  disease  of  the  tube  has  been  present  in 
31  per  cent,  of  our  autopsies.  The  reports  of  other 
gardens  would  indicate  that  their  figures  might  be  quite 
close  to  this.  What  is  the  cause  of  this  high  mortality! 
Incorrect  feeding,  qualitatively  or  quantitatively  has  been 
put  at  the  top  of  the  list  by  Plimmer,  but  he  adds  other 
less  important  factors:  Bacteria  of  infectious  power, 
protozoa,  foreign  bodies  and  parasites  or  their  mural 
cysts.  In  order  to  evaluate  approximately  how  each  of 
these  acts  let  us  review  the  causes  as  they  are  generally 
known  and  later  discuss  the  pathology  as  seen  in  the 
various  orders. 

(1)  Overloading  of  the  stomach  by  too  much  food  or 
by  rapid  eating  of  a  hungry  animal  is  of  importance  under 
certain  domestic  circumstances  where  times  of  feeding 
are  irregular  or  intervals  are  too  long,  but  this  cannot 
occur  in  any  well  regulated  menagerie^  It  is  possible, 
however,  that  overfilling  might  occur  incertain  Ungulata, 
which  have  hay  and  straw  nearly  always  available,  if  the 
food  in  their  reach  happens  to  be  particularly  agreeable 
or  tasty  to  them. 

(2)  Insufficient  mastication  would  seem  to  be  impor- 
tant only  in  those  orders  which  depend  upon  this  action  to 
triturate,  insalivate  and  macerate  their  food,  of  which 
Homo,  Primates,  Ungulata  and  Marsupialia  are  the 
principal  ones. 

(3)  Disturbance  during  and  after  feeding  has  always 
been  believed  to  affect  digestion  unfavorably,  and  it  may 
be  that  visitors  to  a  collection  exert  such  an  effect ;  this 
factor  is  probably  negligible.  ) 


178   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

(4)  The  appropriateness  of  the  food  is  a  very  impor- 
tant factor  in  the  health  of  an  animal  under  captive 
conditions.  Diet  lists  are  made  up  by  officials  largely 
according  to  the  known  habits  and  general  physiology  of 
an  animal,  but  the  food  offered  can  at  best  only  approxi- 
mate what  the  wild  beast  obtains  for  himself.  It  does  not 
follow  because  a  selected  diet  may  seem  to  provide  all  the 
elements  contained  in  the  food  available  under  natural 
conditions  that  it  actually  does  so,  especially  since  we  are 
aware  that  some  essential  food  factors,  known  under  the 
term  vitamins,  are  necessary  to  best  development.  These 
substances  vary  in  closely  similar  foods,  and  seem  to  be 
higher  in  simple  natural  foods  than  in  prepared  diets. 
We  have  seen  in  this  Garden  that  the  inorganic  constitu- 
ents must  be  correctly  represented  in  the  food,  else 
degenerative  osseous  condition  may  develop.  Inappro- 
priate diet  may  express  itself  at  once  after  the  receipt  of 
an  animal,  by  its  sickness  or  death,  or  after  some  time  in 
the  development  of  chronic  tympanites,  chronic  intestinal 
catarrh  or  bony  deformities. 

(5)  The  physical  condition  of  food  is  a  matter  of  no 
small  moment.  The  taking  of  soft  food  in  large  quanti- 
ties especially  by  herbivorous  animals,  permits  too  short 
a  sojourn  in  the  gastric  fundus  and  is  often  followed  by 
pyloric  and  duodenal  disease.  Too  firm  food  may  pack 
the  rumen,  fundus  or  proventricle  as  the  case  may  be,  and 
be  succeeded  by  distention  of  these  parts  and  catarrh  of 
the  pyloric  and  intestinal  area.  The  effect  of  foreign 
bodies  mixed  with  food  is  difficult  to  evaluate  unless  of 
course  they  be  of  such  a  nature  (pointed  metal  and  the 
like)  as  directly  to  traumatize  the  mucosa.  Many  birds 
and  mammals  come  to  autopsy  with  a  relatively  large 
number  of  stones  and  small  sticks  in  the  stomach  without 
any  distinct  evidence  that  they  have  been  hurt  thereby. 
In  the  bird  the  stones  may  be  so  large  and  numerous  as  to 
leave  little  room  for  food,  or  small  enough  to  pass  out  into 
the  intestine  where  they  undoubtedly  may  pave  the  way 


THE  ALIMENTARY  TRACT  179 

for  bacterial  action.  Smoothly  polished  pebbles  in  small 
quantity  seem  to  have  little  effect  in  mammals.  Hair  balls 
are  not  common  and  unless  of  large  size  are  apparently 
unimportant.  Considerable  sand  mixed  with  food  has  a 
distinctly  irritating  effect.  It  is  perhaps  best  kno^vn  as  a 
chronic  gastric  disorder  of  horses;  we  have  seen  it 
in  zebra. 

(6)  Spoiled  food  is  obviously  a  very  prime  factor  in 
inflammations  of  the  gastrointestinal  tract.  Its  opera- 
tions are  illy  understood  except  perhaps  when  products 
of  fermentation  or  putrefaction  prevent  digestive  action 
or  are  absorbed.  If  in  small  quantities  not  sufficient  to 
cause  acute  fermentative  inflammation  or  intoxication, 
such  substances  frequently  taken  may  doubtless  produce 
chronic  catarrhs.  Many  animals  are  fed  upon  vegetable 
mashes,  or  stews  which  can  decompose,  while  bad  meat 
may  occasionally  be  fed.  We  had  a  rather  serious  out- 
break of  enteritis  in  small  Carnivora  from  the  use  of  fowl 
heads  obtained  at  hotels ;  some  of  these  cases  were  shown 
to  be  due  to  B.  paracoli,  thus  to  be  looked  upon  as  infec- 
tions. Dirty  food  while  not  spoiled  may  carry  with  it 
organisms  of  decomposition,  or  of  infective  qualities,  or 
the  dirt  may  act  as  an  irritative  foreign  body.  We  have 
found  that  for  delicate  ungulates  (antelopes)  it  is  highly 
desirable  to  screen  grain,  and  that  the  grade  of  hay 
should  be  of  the  best. 

((7)  Infectious  conditions  are  of  great  importance 
under  certain  circumstances  but  with  the  exception  of  hog 
and  fowl  cholera,  the  dysenteries  and  a  few  other  dis- 
eases, do  not  as  a  rule  play  a  great  part  in  mortality  as 
specific  diseases  unless  of  course  an  epizootic  appear.  The 
greater  problem  is  to  understand  bacterial  action  in  the 
face  of  other  factors.  Are  infectious  germs  introduced 
with  food  and  drink  in  every  case  of  gastroenteritis  or 
do  some  other  factors  activate  those  already  present  in 
the  gut  tract?  Unfortunately  these  questions  cannot  be 
answered  directly.     We  can,  however,  point  out  which 


180   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

groups  of  bacteria  are  most  common  in  some  of  the 
orders,  which  orders  are  most  susceptible  to  bacterial 
invasion  and  which  to  local  lesions  with  intoxication.  The 
greatest  problem  in  the  field  is  the  interrelation  of  germs 
of  various  sorts  in  the  intestinal  tract.  Certain  varieties 
are  known  to  develop  intoxicating  aromatic  substances, 
others  to  elaborate  or  excrete  fatty  acids,  still  others  to 
form  antiferments  but  the  conditions  existing  in  the 
various  kinds  of  intestinal  tracts  are  too  little  understood 
to  help  very  much  in  this  study. 

(8)  Am'mal  parasites  have  long  been  considered  as 
one  of  the  causes  of  gastrointestinal  inflammation,  a  con- 
dition largely  due  to  copying  from  book  to  book  of  a  few 
facts  and  more  impressions.  The  sum  of  reliable  infor- 
mation today  would  seem  to  indicate  that  a  few  parasites 
— uncinaria  being  the  most  conspicuous  example  of  this 
type-t-draw  considerable  blood  from  the  mucosa,  that  a 
few,  like  uncinaria  and  dibothriocephalus^elaborate  an 
absorbable  toxin,  that  some,  notably  ascarids^produce 
an  irritating  substance,  and  tha|fmany  possess  the  power 
in  themselves  or  by  some  excretion  to  act  as  antiferments. 
These  factors,  were  they  all  combined  in  one  worm,  might 
probably  irritate  the  mucosa  sufficiently  to  produce 
^Inflammation,  but  it  is  not  easy  to  imagine  that  thej^  would 
cause  an  acute  specific  condition.  It  is  much  more  easily 
conceived  that  with  tiny  hemorrhages  or  ulcerations  of 
mucosa?,  bacteria  might  get  in  their  work  or  if  consider- 
able ferment  were  neutralized,  maldigestion,  flatulence  or 
indigestive  irritation  would  ensue.  "With  certain  worms 
like  esophagostomum  there  is  considerable  evidence  to 
show  that  a  chronic  fibrous  disease  of  the  intestinal  wall 
arises,  but  in  this  case  the  parasite  resides  in  the  mucosa 
and  acts  as  a  foreign  body.  It  would  seem,  however,  that 
the  most  important  influence  that  animal  parasites  exert 
is  to  be  found  in  the  preparation  of  the  mucosa  for  the 
action  of  bacteria.    Masses  of  parasites  may  of  course 


THE  ALIMENTARY  TRACT  181 

physically  obstruct  the  lumen  and  lead  to  intestinal  stasis 
and  dilatation^) 

Inflammation. 

In  analyzing  the  cause  of  a  gastroenteritis  and  its 
consequent  effect  upon  the  wall  of  the  tube  and  upon  the 
viscera,  certain  physical,  chemical  and  physiological  fac- 
tors must  be  considered.  Whether  this  may  be  directly 
the  effect  of  bacteria  or  poisons  from  worms  or  some 
other  factor  seems  of  little  moment  since  in  any  fully 
developed  case,  symptoms  and  effects  are  comparable. 
Moreover  it  seems  that  pathological  anatomy,  both  gross 
and  minute,  is  essentially  the  same  from  Primates  to 
struthious  birds,  the  highest  and  lowest  of  the  two  classes 
here  considered.  By  this  is  meant  that  the  acute  con- 
gestive condition  of  the  gut  tract  with  solution  of  the 
surface,  to  which  we  have  applied  the  name  of  toxic 
enteritis,  seems  to  be  met  with  in  this  form  throughout 
all  the  orders.  So  too  catarrhal  inflammations  are  the 
same  to  the  naked  eye  and  under  the  microscope,  due 
allowance  being  made  for  the  fact  that  mammals  use  poly- 
nuclear  cells  for  exudative  purposes  while  birds  employ 
mononuclears.  Concomitantly  with  these  conditions,  a 
degenerative  process  may  be  going  on  in  the  liver  and 
kidney,  and  hyperplasias,  especially  in  the  true  infec- 
tive processes,  will  be  found  in  the  related  lym- 
phatic structures. 

The  majority  of  students  today  place  responsibility 
for  gastroenteritis  upon  the  bacteria  known  to  be  present 
in  the  various  intestinal  tracts,  mentioning  especially 
colon  and  proteus  groups,  streptococci,  the  necrosis  bacil- 
lus and  anaerobes  of  the  Welch  class.  In  a  few  of  our 
studies  of  intestinal  bacteria  in  cases  of  enteritis  one  thing 
has  been  very  definite  and  that  is  that  in  the  intestinal 
content  of  animals  whose  food  is  largely  meat,  Gram-nega- 
tive bacilli  have  predominated,  whereas  in  herbivorous 
animals  Gram-positive  organisms  have  been  most  numer- 


182   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 


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184   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

ous.  From  the  observations  of  Kitt,  Strassberger 
and  some  others,  the  normal  flora  of  domesticated  ani- 
mals is  subject  to  wide  variations  so  that  our  observations 
must  receive  confirmation  before  they  are  finally 
acceptable.  We  have  on  several  occasions  isolated  from 
carnivorous  intestines  Bact.  paracoli,  Bact.  suipestifer 
and  other  members  of  this  group.  We  have  no  reliable 
cultural  data  upon  the  herbivorous  intestine  and  can  only 
quote  the  Gram  pictures  as  mentioned  above.  On  two 
occasions,  an  eland  and  an  elk,  a  very  large  number  of 
forms  corresponding  to  necrosis  bacillus  were  seen;  to 
this  organism  Kitt  gives  considerable  power  in  the 
production  of  necrotizing  processes. 

The  Table. 

Our  records  have  been  analyzed  from  the  standpoint 
of  diagnosis  and  the  most  probable  cause.  The  first  will 
be  taken  up  in  discussing  each  of  the  orders.  The  prob- 
able causes  are  divided  into  food,  bacteria,  animal 
parasites,  physical  objects  and  undecided,  in  other  words 
a  classification  based  upon  the  most  prominent  or  definite 
evidences  as  seen  at  postmortem  combined  when  possible 
with  antemortem  observations^  When  findings  were 
inconclusive  or  contradictory,  cases  were  called  unde- 
cided, naturally  a  very  large  group.  Fermentative 
processes  in  the  presence  of  obviously  undigestible 
material,  are  classified  with  food  as  a  cause.  When  evi- 
dences of  septicemia  existed  in  absence  of  the  other 
factors,  it  is  held  that  bacteria  were  responsible.  Cases 
were  grouped  under  animal  parasites  when  these  were  the 
most  definite  findings.  Physical  objects  are  relatively 
unimportant  and  self-explanatory.  To  the  etiological 
chart  there  are  appended  columns  intended  to  show  the 
percentage  or  case  incidence  of  the  disease  of  the  grosser 
subdivisions  of  the  gastrointestinal  tract  which  indicate 
in  a  general  manner  what  part  of  the  tube  in  the  various 
orders  is  most  susceptible  to  disease.    While  of  course 


THE  ALIMENTARY  TRACT  185 

conclusions  must  be  drawn  with  great  caution,  there  can 
be  little  doubt,  for  example,  that  carnivores  and  mar- 
supials have  more  gastric  disease  than  any  other  order, 
and  that  the  high  place  for  the  colon  is  held  by  the  galli- 
naceous birds.  This  charting  was  suggested  by  the  work 
of  Dr.  Raymond  Pearl  upon  statistics,  wherein  he  takes 
as  a  basis  of  classification  the  part  of  the  body  which 
succumbs  to  disease-producing  organisms  or  from  which 
a  disease  starts.  It  caimot  be  stated  that  there  is  a  clear 
cut  relationship  between  enteritis  and  the  expectancy 
of  life. 

Mammalia. 

The  Primates  as  an  order  have  their  share  of  in- 
flammations of  the  gastrointestinal  tract  and  present 
points  of  interest.  Acute  digestive  disorders  succeeded  by 
acute  dilatation  of  the  stomach,  or  in  less  fermentative 
cases  by  acute  catarrh  of  the  intestine,  are  not  at  all  un- 
common. The  reason  for  this  is  not  discovered  by  review- 
ing the  diet  and  manner  of  feeding.  The  buccal  pouches, 
distensible  esophagus,  the  freely  movable  stomach,  and 
relatively  elastic  gastric  wall  would  seem  to  permit  of  very 
considerable  dilatation  to  accommodate  the  large  quanti- 
ties which  the  monkey  sometimes  crams  into  himself. 
Nine  fairly  acceptable  records  of  gastric  overfilling  exist 
and  two  of  them  seem  to  have  been  followed  by  tympan- 
ites sufficient  to  embarrass  respiration,  in  one  case  there 
occurring  an  acute  cardiac  dilatation  with  myocardial 
degeneration.  The  animals  give  no  symptoms  of  this 
condition  and  in  the  last  case  cited  the  beast,  while  old, 
ate  well  and  was  not  distended  the  evening  before  death. 

When  acute  gastritis  exists  (twenty  cases)  the  animal 
seems  uneasy  but  does  not  vomit.  On  one  occasion  I  was 
called  to  see  a  monkey  which  was  retching  and  seemed  in 
pain.  Lime  juice  was  offered  and  taken,  followed  by 
gentian  and  cardamon,  which  seemed  to  give  some  bene- 

13 


186  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

fit.  Somewhat  later  this  was  repeated  in  another  case, 
but  observations  where  this  might  be  useful  are  rare. 

The  pathology  of  gastric  conditions  offers  little  to 
contrast  with  that  of  man.  The  enormous  distensibility 
of  the  fundal  pouch  often  suggests  to  the  observer  the 
rumen  of  ungulates.  Acute  gastritis  of  one  kind  or 
another  and  acute  catarrhal  enteritis  are  the  most  com- 
mon lesions  noted  in  the  Primates.  Involvement  of  the 
intestine  or  colon  need  not  carry  with  it  an  increase  of 
signs  of  illness,  although  at  times  one  will  see  an  evi- 
dently sick  animal  with  diarrhcEa.  Anatomically  the 
lesions  are  commonly  restricted  to  the  stretches  of  gut 
above  the  ileum,  it  being  rather  rare  that  this  division  or 
the  colon  is  affected.  Pathologically  the  lesions  are 
catarrhal  with  definite  involvement  of  the  follicles  in 
about  one-third  of  the  cases.  In  this  order  toxic  and 
pseudomembranous  forms  are  quite  rare  and  ulcerative 
lesions  uncommon.  Colonic  disease  as  a  sequel  to  inflam- 
mation higher  up  is  sometimes  seen  in  the  follicular 
varieties,  but  takes  a  minor  place  compared  to  amoebic 
dysentery  of  which  we  have  had  several  cases ;  this  will 
be  discussed  under  a  separate  heading.  Degenerative 
disease  of  the  skeleton  is  almost  always  accompanied  by 
a  low  grade  of  enteritis  but  not  necessarily  gastritis  or 
colitis.  The  pallor  of  the  mucosa,  while  at  times  striking, 
may  be  relieved  by  follicular  spots  and  petechia  or  pig- 
mentation. Often,  however,  animals  suffering  from 
osteomalacia  and  rickets  come  to  their  end  by  an  acute 
inflammation  of  the  gut  tract. 

The  bacteriology  at  our  disposal  allows  no  conclusions. 
Aside  from  a  case  probably  due  to  Ps.  fluorescens  and 
one  with  colon  bacillus  abscesses  in  the  liver,  no  reliable 
data  are  at  hand. 

Reference  to  Table  11  reveals  the  fact  that  among 
orders  with  sufficient  autopsies  to  permit  percentages,  the 
alimentary  tract  in  monkeys  is  in  the  group  of  low  figures, 
that    the    intestinal    section    is    relatively    more    often 


THE  ALIMENTARY  TRACT  187 

affected,  and  that  the  colon  is  more  often  diseased  than 
in  other  mammalian  orders,  and  is  exceeded  only  by  the 
gallinaceous  birds. 

The  Lemures,  of  which  we  have  eighty-six  autop- 
sies, do  not  differ  much  in  anatomy  from  the  Primates, 
however  greatly  they  disagree  in  habits  and  outward 
appearance ;  their  diet  is  the  same.  Clinically  the  sloth- 
ful behavior  of  a  normal  lemur  probably  obscures 
symptoms  and  signs  of  illness,  for  our  antemortem  notes 
with  the  exception  of  a  few  observations  of  loose  stools, 
fail  to  offer  a  lead  as  to  diagnosis.  This  order  has  a  large 
incidence  (twenty-three  cases)  of  gastroenteric  condi- 
tions as  shown  in  Table  11,  but  some  explanation  of  the 
figures  is  deserved.  In  the  first  place,  only  one  case  of 
acute  gastritis  occurred,  and  this  was  apparently  a  part 
of  a  general  infection,  and  if  induced  at  all  by  food  this 
was  only  secondary.  Indeed  as  one  reviews  the  records 
it  does  not  seem  that  the  lemurs  are  easily  disturbed  in 
their  gastric  digestion.  Acute  and  subacute  inflamma- 
tions from  bacterial  action  seem  definitely  more  promi- 
nent since  they  take  the  catarrhal,  follicular  and  deep 
submucous  form  and  are  frequently  associated  with  gen- 
eralized infectious  processes.  One  amoebic  case  was 
observed  and  there  was  another  in  which  a  heavy  cestode 
and  nematode  infestation  seemed  to  have  paved  the  way 
for  bacteria. 

Carnivora. 

The  food  of  this  order  is  received  into  the  fundal 
part  of  the  stomach,  the  distensible  but  normally  capa- 
cious left  and  superior  two-thirds  of  the  organ.  The 
general  shape  of  the  viscus,  that  of  a  gourd,  permits  a 
fairly  sharp  separation  of  the  fundal  and  pyloric  sections, 
so  definite  indeed  that  the  pathology  of  the  two  parts  was 
studied.  The  intestines  vary  in  length,  but  in  the 
land  carnivores  are  relatively  short,  narrow  in  lumen 
and  rich  in  waU.    A  cecum,  or  at  least  a  blind  end  of  the 


188  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

large  gut  made  by  the  insertion  of  the  small  intestine 
above  the  tip  of  the  colon,  is  suggested  in  all  families, 
although,  as  in  the  bears,  it  may  be  quite  insignificant  or 
rudimentary.  Theoretically  no  stasis  should  occur  at  this 
point.  The  colon  is  short  in  all  carnivores  and,  like  the 
small  gut,  with  a  heavy  wall.  The  comparative  simphcity 
of  the  carnivorous  gut  tract,  the  ability  of  many  of  these 
animals  to  disgorge,  the  suggestion  of  high  resistance  of 
the  upper  end  of  the  tract  to  infection  and  the  ease  with 
which  diarrhoea  can  clear  out  the  tube,  would  seem  to 
warrant  the  expectation  that  inflammation  would  not  be 
serious.  Such,  however,  is  not  the  fact  for,  on  the  con- 
trary, they  have  shown  a  higher  incidence  than  any  other 
order  for  which  we  have  adequate  comparison.  Anatomi- 
cally considered  their  stomach  occupies  the  second  place 
in  vulnerability,  next  to  the  marsupials,  and  their 
intestines  the  highest  place ;  this  indicates  of  course  that 
combined  gastric  and  intestinal  disease  has  often  occur- 
red. Involvement  of  the  colon  occupies  the  second  place, 
in  ordinate  susceptibility,  being  exceeded  only  by  the 
monkeys,  due  to  heavy  parasitic  infestation,  but  would 
occupy  the  first  place  were  the  eleven  amoebic  dysenteries 
in  monkeys  subtracted  from  their  total,  a  subtraction 
which  might  be  allowed  since  it  represented  an  epi- 
zootic outbreak. 

Etiologically  considered,  it  would  seem  as  if  the 
influence  of  incorrect  feeding  were  of  little  importance, 
and  from  one  standpoint  this  is  probably  the  case.  Acute 
fermentative  or  irritative  processes  are  not  common  at 
all,  while  more  inflammatory  pictures,  catarrhal,  erosive 
or  ulcerative,  are  the  rule.  There  is  another  phase  to  the 
term  incorrect  food,  that  is  incorrect  in  its  cleanness. 
During  1912-15  there  was  an  increasing  mortality  among 
the  cats  and  dogs  fed  upon  horse  meat,  mutton  and  fowl 
heads.  Early  in  1916  the  butcher  shop  was  reconstructed 
and  thoroughly  cleaned  and  covered  galvanized  pans 
supplied  in  which  to  transport  the  food ;  these  pans  were 


THE  ALIMENTARY  TRACT  189 

scrubbed  and  scalded  after  use.  Since  that  time,  infec- 
tious inflammations  of  the  stomach  and  intestines  have 
shown  an  ever  increasing  downward  incidence,  which 
result,  there  having  been  no  material  changes  in  other 
directions,  I  do  not  hesitate  to  ascribe  to  the  improve- 
ment of  butchering  and  dispensing  engineered  by  Dr. 
W.  B.  Cadwalader. 

Helminths  seem  to  be  of  importance  in  this  order  both 
by  reason  of  the  percentage  of  autopsies  in  which  they 
presented  the  most  probable  or  at  least  most  suggestive 
cause  and  because  uncinaria  and  strongylus  have  been 
seen  attached  to  the  wall  and  a  large  bulk  of  known  irri- 
tative cestodes  have  occupied  the  lumen.  Physical 
objects,  stones,  bones,  wire,  may  cause  irritation  enough 
to  activate  bacterial  action  or  may  actually  penetrate  the 
wall;  the  latter  action  is  well  known.  In  so  far  as  prac- 
tical application  of  this  is  concerned,  it  teaches  to  feed 
whole,  unsplintered  or  ground  bone. 

The  distribution  and  character  of  pathological  lesions 
according  to  the  region  of  the  stomach  is  what  might  be 
expected  from  the  shape  and  physiology  of  its  parts. 
True  inflammatory  processes  are  best,  and  in  some  cases 
only  seen  in  the  pyloric  half  of  the  viscus,  while  the 
changes  in  those  few  cases  believed  to  be  fermentative  or 
irritative  in  nature  were  largely  confined  to  the  fundus. 
Dilatation  of  the  latter  part  may  be  understood  because 
there  the  muscular  coats  are  about  equal  to  the  mucous  in 
thickness  and  one-half  the  width  of  those  at  the  pylorus, 
but  why  inflammatory  processes  should  not  be  so 
developed  in  the  fundus  is  not  clear  unless  the  greater 
availability  of  mucus  protects  the  secreting  wall.  Not 
only  does  acute  inflammation  reach  its  most  definite 
form  in  the  second  part  of  the  stomach,  but  the  irregular 
pigmentation,  mammillated  overgrowth  and  atrophy  or 
ulceration  of  chronic  disease  are  likewise  best  seen  in 
this  part. 


190  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Acute  enteritis,  of  all  varieties,  is  seen  more  beauti- 
fully in  carnivores  than  in  any  other  order  of  mammals, 
and  nowhere  can  it  be  studied  better.  Its  gross  appear- 
ance is  that  of  the  text-book  and  its  minute  character  even 
more  instructive.  I  have  used  a  slide  of  acute  catarrhal 
enteritis  in  a  lion  for  the  illustration  of  this  lesion  for  the 
Text-hook  of  Pathology  by  Doctor  Stengel  and  myself. 
(However,  as  is  known  to  all  who  have  paid  any  attention 
to  enteritis,  the  postmortem  findings  are  usually  much 
less  definite  than  clinical  observations  would  warrant  one 
to  expect.  The  Carnivora  not  uncommonly  show  intestinal 
congestion,  mucous  membrane  swelling  without  edema 
or  opacity,  congestion  of  the  spleen,  cloudy  swelling  of 
the  liver  and  kidneys  and  perhaps  mesenteric  lymph  node 
edema.  This  picture  we  have  viewed  as  a  toxic  affair  of 
some  sort  or  a  bacterial  infection  not  yet  far  enough 
advanced  to  produce  catarrhal  or  ulcerative  enteritis 
and  septicemia.  In  such  cases  the  carnivorous  intestinal 
mucosa  offers  instruction.  The  epithelium  is  vacuolated 
or  fringed  on  the  free  edge  or  may  be  missing  altogether. 
In  the  depths  mucus  formation  is  very  active,  and  where 
it  is  going  on,  round  cells  seem  attracted,  collecting  in 
groups  in  the  villus  or  in  the  subjacent  submucosa.  Peri- 
vascular round  cell  increase  may  be  noted.  Plasma  cells 
and  granular  eosinophiles  are  common,  but  I  cannot  state 
how  important  the  latter  are  in  the  general  picture 
because  of  the  frequency  of  parasites  in  carnivores.  The 
central  vessel  of  the  villus  and  the  arterioles  of  the  sub- 
mucosa are  injected.  Lymph  follicles  may  or  may  not  be 
enlarged,  but  if  so  usually  fail  to  show  a  germ  centre. 

Colitis  alone  is  not  common  in  this  order,  but  as  an 
extension  process  or  involvement  at  the  same  time  as  the 
upper  levels  it  occurs  occasionally.  The  only  fact  I  wish 
to  record  and  one  which  I  would  emphasize  because  of 
having  seen  it  recently  in  a  human  case  of  chronic  colitis, 
and  since  it  does  not  appear  important  to  systematic 
writers,  is  superficial  blood  supply.     The  capillary  net- 


THE  ALIMENTARY  TRACT  191 

work  of  the  colonic  villi,  while  rich,  is  in  the  form  of  a 
fine  plexus  just  under  the  epithelium.  In  the  cases  studied 
these  vessels  become  quite  distinct  and  possess  much 
more  definite  walls,  often  bordered  by  mononuclears,  while 
connective  tissue  is  more  evident  at  the  bases  of  the  villi 
and  deeper.  This  may  help  in  deciding  the  existence  of 
a  colitis. 

Bacteriologically  the  most  instructive  experience  to 
report  is  the  discovery  that  a  small  outbreak  of  enteritis 
among  small  Carnivora,  chiefly  cats,  fed  upon  fowl  heads 
was  due  to  Bact.  paracoli,  or  at  least  this  organism  was 
found  in  the  intestinal  mucosa,  spleen,  and  heart's  blood 
of  three  cases.  The  type  of  enteritis  was  hemorrhagic  and 
follicular.  There  was  also  a  case  of  septicemia  appar- 
ently emanating  from  enteritis  due  to  Bact.  suipestif er  in 
a  lion  {Felis  leo).  These  facts  bring  strongly  to  attention 
the  modem  teaching  that  meat  poisonings  of  the  Gartner 
type  are  to  be  considered  as  infectious  and  not  of  the 
so-called  ptomaine  group. 

PiNNiPEDiA,  while  related  closely  to  the  Carnivora, 
are  grouped  in  a  suborder  in  our  classification  and  because 
of  their  restricted  diet  are  treated  here  in  a  separate 
paragraph.  The  tract  is  peculiar  in  the  strong  tubular 
stomach  sharply  bent  upon  itself,  the  great  length  of  the 
small  gut  (upwards  of  a  hundred  feet  in  some  genera), 
and  the  practical  absence  of  a  cecum.  Pathologically 
speaking,  the  most  striking  lesion  of  these  animals  is 
ulcerative  gastritis,  a  process  usually  most  marked  along 
the  posterior-superior  surface,  but  not  confined  thereto. 
Upon  inspection  the  gastric  mucosa,  normally  supplied 
with  low  regular  rugae,  is  much  distorted  by  swellings 
upon  the  top  of  which  are  irregular  ragged  ulcers  with 
rounded  elevated  but  not  frayed  margins.  The  density 
of  the  edges  indicates  much  infiltration  of  the  deep 
mucosa  and  submucosa;  this  can  be  confirmed  by  micro- 
scopical examination.  One  attempt  to  study  this  gastritis 
bacteriologically  was   fruitless.     Sections   of  one   case 


192   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

showed  streptothrix-like  masses  while  in  another  case 
bacterial  colonies  and  yeast-like  bodies  were  found  in 
adjacent  lymph  nodes.  The  genesis  of  this  condition 
might  lie  in  injury  by  fish  fins  or  by  foreign  bodies,  of 
which  large  numbers  are  found  at  times  (a  pint  and  a  half 
of  stones,  marbles,  and  sticks  were  found  in  one  stomach). 
fGastritis  has  been  the  starting  point  of  septicemia  on  two 
occasions,  and  three  times  an  acute  exacerbation  or  new 
implantation  of  infection  occurred,  ^vith  extension  into 
the  intestine.  It  is  interesting  that  all  the  deaths  of 
Pinnipedia  with  gastroenteric  conditions  occurred  in  the 
winter  months. 

Insectivora  are  represented  by  two  common  Euro- 
pean Hedgehogs.  In  one  there  were  three  shallow 
but  shelving  ulcers  in  the  stomach  which  had  bled  suffi- 
ciently to  weaken  the  animal;  free  blood  was  found  in  the 
intestine.  The  other  specimen  was  diagnosed  at  post- 
mortem as  having  catarrhal  enteritis  involving  nearly 
the  whole  small  gut,  but  histological  section  did  not  con- 
firm this. 

As  one  descends  the  zoological  scale  the  first  gastro- 
intestinal tract  prepared  for  the  nutritional  care  of  bulky 
food  is  to  be  found  in  the  Rodentia.  This  order  pre- 
sents a  great  variety  of  shapes  and  arrangements  of  the 
stomach,  but  the  outstanding  feature,  with  very  few 
exceptions  (cf.  spermophiles),  is  the  development  of  the 
cardiac  and  fundal  divisions  ostensibly  for  the  reception 
of  a  large  bulk  of  coarse  food  to  be  digested  at  leisure. 
Some  genera  like  the  hamster  {Cricetus)  have  a  stomach 
closely  resembling  the  ruminants,  while  that  of  the 
spermophile  suggests  the  equine  stomach.  The  pyloric 
end,  variable  in  many  ways,  greatly  resembles  the 
abomasum.  So  too  the  duodenum  is  large,  loose  and  dis- 
tensible while  the  copious  small  gut  ends  in  a  very  large 
cecum,  shaped  at  times  in  a  manner  which  has  led  to  the 
term  ''colonic  stomach."  The  colon  is  variable  and  not 
always  supplied  with  longitudinal  bands  and  sacculations. 


THE  ALIMENTARY  TRACT  193 

In  regard  to  incidence  of  gastrointestinal  disease, 
rodents  occupy  a  middle  position  in  the  table.  The 
stomach  seems  a  vulnerable  section  of  the  tract.  Dilata- 
tion of  the  left  hand  section  is  common,  due,  to  all  appear- 
ances, to  fermentative  processes  which  have  as  a  result 
the  softening  of  the  mucosa  so  that  even  immediately- 
after  death  it  will  separate  almost  entirely.  In  these 
cases  the  pyloric  part  need  not  participate  but  may 
remain  flat,  smooth,  soft  and  pink.  This  condition  is 
slightly  more  common  in  the  compound  than  in  the  simple 
stomachs.  In  some  of  this  order,  especially  rats  and 
cavies,  there  is  a  fermentative  gastroenteritis  expressed 
by  injection  and  edema  of  the  pylorus  and  duodenum,  and 
much  frothy  mucus.  It  was  at  first  thought  that  some 
relation  might  exist  between  this  condition  and  the 
absence  of  the  gall-bladder,  but  it  occurs  in  varieties 
possessing  this  structure.  The  reaction  of  the  intestine 
to  irritation  in  this  order  is  peculiar  in  two  ways,  the 
occurrence  of  mucus  and  the  activity  of  the  lymphatics. 
In  all  the  inflammations  from  and  including  the  stomach 
to  the  cecum,  mucus  is  conspicuous.  At  times  it  is  thin 
or  loose  and  mixed  with  contents,  while  at  others  it  forms 
a  relatively  close  covering  for  the  mucosa  almost  like  a 
false  membrane.  Rodentia  are  peculiar  in  the  prompt- 
ness and  clearness  with  which  the  follicles  of  the  intestinal 
wall  and  mesentery  enlarge  in  inflammation.  They 
appear  as  pale,  well  outlined  or  diffuse  opacities  in  the 
wall  or  as  distinct  plaques  prominent  on  the  surface. 

The  Proboscidea  are  represented  by  one  Elephant 
{Elephas  indicus),  in  which  a  mild  catarrhal  change 
was  seen  in  the  middle  stretches  of  the  small  intestine. 
This  was  of  little  importance  as  a  cause  of  death,  there 
being  several  other  diagnoses,  and  was  probably  a  termi- 
nal affair. 

Hyraces,  of  which  we  have  a  total  of  seven  ex- 
amples, present  two  mild  involvements  of  the  intestine  but 
none  of  the  stomach.    It  would  appear  from  the  records 


194  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

that  the  intestinal  condition  had  little  to  do  with  the  death 
of  the  animals,  and  unfortunately  no  microscopic  slides 
were  made.  Because  of  the  curious  formation  of  the  large 
gut,  notes  of  both  ceca  were  made  in  one  case,  and  can  be 
condensed  as  follows:  "The  upper  or  anterior  cecum 
presented  a  shaggy  pearl  gray  mucous  covering,  closely 
attached  to  the  mucosa.  This  cavity  and  the  posterior 
ceca  were  packed  with  dry  crumbling  feces.  Small  thin- 
walled  cysts  were  seen  in  the  tips  of  the  lower  ceca.  Duo- 
denum was  congested  and  mucosa  swollen.  Stomach 
contained  dry,  poorly  digested  food.  No  parasites 
were  found. ' ' 

The  Edentata  are  represented  by  an  Armadillo 
{Tatu  novemcinctiis)  and  an  Ant-eater  {Myrmecophaga 
tridactyla.)  The  former  had  a  prolapse  of  the  rectum 
accompanied  by  colitis  but  it  is  impossible  to  decide  the 
priority  of  the  two  conditions  since  the  former  is  known 
to  have  existed  long  enough  to  have  permitted  the  latter 
to  develop.  The  Ant-eater  had  a  distinct  mucocatarrhal 
enteritis  in  which  bacteria  played  a  part  since  involve- 
ment of  the  liver,  spleen,  kidney  and  lymph  nodes  also 
existed.  The  beast  was  in  good  condition  upon  arrival, 
but  did  not  become  accustomed  to  the  proffered  diet,  and 
was  distinctly  anemic  at  death. 

The  Ungulata,  so-called  for  their  hoofed  and 
horned  character,  are  also  associated  anatomically  by  the 
construction  of  their  gastrointestinal  tract.  However,  the 
order  of  the  list  as  given  on  page  44  does  not  represent 
their  historical  development  nor  does  it  accord  with 
anatomical  arrangement  of  the  tube  under  discussion. 
The  last  three  families  of  Artiodactyla  are  perhaps  the 
simplest  in  the  construction  of  this  tube,  or  at  least  take 
an  equal  place  mth  the  Peris sodactyla,  while  the  remain- 
ing families  of  the  former  suborder  have  a  complex  tract 
of  generally  similar  architecture.  Tliis  whole  order  has, 
however,  an  alimentary  tract  anatomically  suited  for  the 
consumption  of  bulky  vegetable   stuffs   and   shows   an 


THE  ALIMENTARY  TRACT  195 

attempt  at  adjustment  between  the  food,  tlie  methods  of 
mastication,  the  area  of  digestive  surface,  and  the  bulk 
necessary  for  nutrition.  The  number  of  factors  opened 
up  by  the  many  variations  between  this  complex  tract  and 
that  of  the  carnivorous  simple  tract  is  so  great  that  I  shall 
attempt  only  to  contrast  the  anatomy  and  pathology  of 
the  simpler  ungulate  tracts  and  the  complex  rumi- 
nant apparatus. 

The  simpler  gastrointestinal  tract  is  possessed  by  the 
Perissodactyla,  and  by  the  follomng  families  of  Artio- 
dactyla,  the  Phachocoeridae,  Suidas  and  TayassuidaB.  This 
consists  of  a  stomach  with  a  single  cavity  (some  Peccaries 
have  partitions  but  no  true  septa  with  strict  histological 
differences)  divided  into  esophageal,  cardiac,  fundal  and 
pyloric  areas,  dependent  upon  the  nature  of  the  lining 
epithelia  and  the  absence  or  presence  of  glands,  as  well  as 
the  nature  of  the  tubules.  The  duodenal  section  is  ample 
and  may  be  sacculated  while  the  intestines  are  small  in 
calibre  and  rather  sturdy  in  wall.  The  cecum  is  rela- 
tively very  large,  well  supplied  by  longitudinal  bands  and 
sacculations;  the  colon  is  relatively  short  but  quite 
capacious  and  sacculated. 

The  ruminants  and  other  remaining  members  of  the 
Artiodactyla  have  a  compound  stomach  suited  to  the 
separation  of  coarse  and  fluid  foods  and  the  retention  of 
water,  and  so  arranged  that  boluses  of  different  densities 
are  distributed  a&  needed.  These  divisions  are  histo- 
logically as  well  as  grossly  different.  The  first  three, 
comparable  to  the  esophageal  section  of  the  simpler 
stomach,  are  reservoirs  or  channels,  while  the  fourth  or 
true  digestive  section,  is  divided  into  areas  corresponding 
to  fundus  and  pylorus,  possessing  the  appropriate  type 
of  gland.  The  duodenum  in  this  group  is  narrow,  as  is 
the  rest  of  the  small  gut,  and  has  delicate  walls.  The 
cecum  proper  is  short  and  of  variable  width,  but  never  as 
great  as  in  the  group  first  discussed,  while  the  colon,  an 
intricately  wound  tube,  is  narrow  and  very  long.    Certain 


196   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

of  the  first  group  (Peccaries  )have  a  colon  of  this  type, 
but  it  is  not  so  complicated  as  in  time  ruminants. 

These  complications  seem  designed  to  permit  of  a  long 
retention  of  coarse  food  of  low  nutritive  value  per  bulk 
for  a  time  sufficient  for  full  digestion ;  reverse  adaptation 
of  large  size  of  stomach  and  colon  may  be  explained  on  the 
same  basis.  Let  us  now  examine  these  two  groups  to  dis- 
cover their  pathological  reactions  and  the  nature  of 
the  lesions. 

Ungulates  as  a  class  stand  in  a  position  equivalent  to 
the  Primates  in  the  incidence  of  gastrointestinal  diseases. 
They  show  a  conspicuous  percentage  of  cases  apparently 
due  to  incorrect  food  and,  coupled  with  this,  a  high  per- 
centage of  gastric  involvement,  being  exceeded  in  this 
respect  by  the  marsupials  only  (there  is  an  adequate 
reason  for  this — see  below).    Cases  ascribed  to  bacterial 

Table  12. 

Showing  the  Percentage  Incidence  of  Various  Forms  of  Inflammation  in  the 
Alimentary  Tract  of  Ungulata. 


Lesion 

Group  A 
Periasodactyla  and 
Swine,  Wart  Hogs, 

Peccaries 

Per  cent. 

Group  R 

Ruminants  and 

Relatives 

Per  cent. 

Acute  fermentative  gastritis 

All  other  forms  of  gastritis 

Acute  toxic  or  fermentative  enter- 
itis  

18. 
16.3 

1.6 
1.6 
0. 
0. 

2.1 
5. 

2.3 

All  other  forms  of  acute  enteritis .  . . 
Chronic  enteritis 

8.4 
1.9 

Cohtis  and  typhlitis 

3.1 

or  parasitic  agents  are  not  numerous.  Analysis  of 
the  records  of  the  two  groups  discussed  above  brings  out 
some  interesting  facts.  Consultation  of  Table  12  shows 
percentage  comparisons,  based  on  the  number  of 
autopsies,  of  lesions  in  the  various  sections  of  the  tract. 
Group  A,  that  with  the  simple  stomach  and  the  short 
capacious  colon,  is  represented  by  forty-four  specimens, 
while  Group  B  embraces  321  animals.  Perhaps  the  use  of 
these  two  widely  differing  figures  for  comparison  is  open 


THE  ALIMENTARY  TRACT  197 

to  some  objection  which  might  be  final  and  decisive  were 
not  the  figures  so  definite.  At  a  glance  one  can  see  that 
Group  A  has  involvement  more  marked  high  in  the  tract 
while  Group  B  has  more  disease  in  the  intestine  and  colon. 
Certainly  gastritis  is  more  common  (five  times)  in  the  A 
than  in  the  B  group,  while  enteritis  is  more  common  in  B 
(over  three  times).  No  case  of  chronic  enteritis  or  of 
involvement  of  the  colon  is  recorded  in  animals  with  a 
simple  stomach  and  a  wide  hind-gut.  This  may  be  read 
either  in  terms  of  vulnerability  of  the  stomach  or  in  the 
degree  of  resistance  of  the  respective  groups. 

Consideration  of  the  local  factors  of  the  stomach 
brings  to  light  at  once  the  fact  that  incorrect  food  enter- 
ing the  simple  stomach  could  attack  the  softer,  less 
resistant  glandular  section  of  the  fundal  and  pyloric 
areas  whereas  the.  rumen  and  psalter  of  the  compound 
organ,  with  their  stratified  epithelium  devoid  of  glands, 
act  as  barriers  or  as  places  where  detoxication  of  irri- 
tants might  take  place.  In  both  groups  bulky  food  is 
packed  to  the  left,  the  esophageal  and  cardiac  section 
in  the  simple  form,  the  rumen  in  the  compound.  Soft  or 
liquid  food  may  pass  into  the  psalter  and  abomasum  of 
the  ruminant  stomach  almost  directly  since  it  has  not  the 
force  or  bulk  to  push  aside  the  valve-like  fold  of  wall  at 
the  junction  of  esophagus,  rumen  and  reticulum.  For 
this  reason,  if  for  no  other,  the  character  of  soft  food 
supplied  to  this  order  must  be  unexceptionable. 

It  has  not  been  possible  to  follow  out  the  layering  of 
diets  as  Scheunert  did  when  showing  the  course  of  various 
foods  before  they  are  mixed  at  the  beginning  of  the 
pyloric  compartments.  We  have  seen  two  cases  in 
ruminants  which  seem  to  indicate  that  soft  food  had 
passed  into  the  right  side  cavities  of  the  stomach,  there  to 
cause  irritation,  while  the  rumen  remained  quite  normal. 
It  seems,  however,  accepted  by  veterinarians  that  exces- 
sive soft  food  may  be  followed  by  trouble  in  the  digestive 
stomachs,  while  excessive  dry  food  may  cause  distention 


198  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

of  the  left  hand  sections.  So  far  as  I  know,  the  relative 
incidence  of  gastric  disorders  in  the  above  outlined 
groups  has  not  been  pointed  out  before. 

The  pathological  types  of  gastroenteritis  do  not  pre- 
sent many  unusual  features.  Simple  injection  of  vessels 
during  digestion  seems  more  evident  in  this  than  any 
other  order,  while  mucus  production  seems  less  marked. 
The  mucosa  of  the  reservoir  portions  may,  in  simple  over- 
filling and  fermentative  distention  (gastric  tympanites), 
be  quite  normal  or  dull  red ;  when  active  fermentation  has 
occurred  it  may  be  digested  and  peel  off.  More  or  less 
true  inflammation  as  seen  in  the  digestive  stomachs  is 
definitely  better  expressed  in  the  compound  ruminant 
organ  than  in  the  simple  equine  variety  where  congestion 
and  edematous  swelling  with  hemorrhage  form  the  usual 
picture.  True  catarrhal  changes  both  gross  and  minute, 
are  often  well  seen  and  ulcerative  lesions  are  not 
uncommon;  ulcers  are  unusual  in  the  simple  stomach. 
These  statements  hold  good  also  for  the  small  intestine. 
Enteritis  affects  the  duodenum  and  jejunum  more  in  the 
Perissodactyla  and  swine  than  in  the  ruminants. 
Ungulata  do  not  react  with  hyperplasia  of  the  mural 
IjTuphatics  as  do  many  other  orders,  but  the  swelling  of 
the  mesenteric  nodes  is  often  noteworthy.  As  might  be 
expected  this  is  more  definite  mth  catarrhal  changes  and 
therefore  best  seen  in  the  ruminants.  In  the  colon,  how- 
ever, the  solitary  follicles  are  often  quite  prominent  in 
simple  inflammation.  Histologically  the  greatest  changes 
seen  in  this  order  are  superficial  degenerations  with 
mononuclear  increase  in  the  deep  submucosa,  mostly 
arranged  in  perivascular  fashion.  The  villi  do  not  con- 
tain the  large  number  of  cells  seen  in  the  Carnivora. 

Maksupialia. 

Consultation  of  the  figures  for  this  order  in 
Table  11  arrests  attention  at  once  because  of  the 
peculiar  percentages  found  under  the  detailed  factors. 


THE  ALIMENTARY  TRACT  199 

Thus  food  is  of  no  moment  as  a  disturber  of  the  alimen- 
tary tract,  while  bacteria  and  parasites  are  high  in 
importance.  The  gastric  segment  is  more  often  attacked 
than  any  other  portion  and  slightly  more  often  than  in  the 
nearest  order,  Camivora.  These  unusual  figures  deserve 
explanation,  to  which  purpose  it  will  be  necessary  to  con- 
sider the  anatomy  of  the  organ  and  to  discuss  why 
bacteria  and  parasites  stand  so  high  in  etiology. 

Marsupials  are  divided  into  six  families  in  our  classi- 
fication (see  page  44)  which  in  regard  to  their  diet, 
range  from  largely  carnivorous  (the  first  two)  through 
those  choosing  mixed  insects,  fruits  and  vegetables  (the 
second  two)  to  those  eating  vegetables  and  grain  (the  last 
two).  The  stomach  of  these  animals  does  not  vary  exactly 
according  to  their  diet,  the  first  four,  opossums,  dasyures, 
bandicoots  and  wombats,  possessing  an  organ  closely 
similar  in  outline  and  construction  and  resembling  the 
carnivorous  variety,  while  the  phalangers  and  kangaroos 
have  a  stomach  entirely  different  from  the  first  four 
although  somewhat  similar  to  one  another.  The  first 
group  has  a  round  or  irregularly  elliptical  organ  with  the 
esophagus  and  pylorus  close  together  along  the  lesser 
curvature.  The  wombats  have  a  bank  of  glands  sur- 
rounded by  a  capsule,  near  the  cardia.  The  stomach  of 
the  first  four  animals  is  divided  into  cardiac,  fundal  and 
pyloric  parts  by  the  construction  of  their  mucosa,  the 
first  mentioned  division  being  a  high,  rounded  pouch, 
rumen-like,  well  to  the  left.  The  phalanger  's  stomach  is 
more  elongated,  the  two  openings  well  separated  and  a 
fissure  is  found  in  the  right  end  of  the  lesser  curvature 
which  serves  to  separate  the  pyloric  part  from  the  rest. 
The  Macropodidffi  all  have  a  stomach  resembling  the 
human  colon  in  being  elongated,  with  longitudinal  bands 
gathering  it  into  sacculations.  There  is  a  distinct  eso- 
phageal section  to  the  left  with  a  blind  sac  suggesting  an 
ungulate  rumen,  a  long  tubular  fundal,  and  a  sacculate 
pyloric  division. 


200   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 


The  small  intestine  of  the  order  starts  with  the  opos- 
sums as  a  stout  muscular  and  mucous  tube  fitted  for  meat 
eating,  but  as  one  proceeds  to  study  the  families  do^vIl- 
ward  in  the  list  this  tube  becomes  more  delicate  and 
longer.  In  the  first  two  families  the  cecum  is  rudimentary 
and  the  colon  very  short  as  in  cats,  but  the  length  and 
capacity  of  these  parts  increase  through  the  bandicoots 
and  wombats  until  in  the  strict  fruit,  vegetable  and 
grain  eaters,  phalangers  and  kangaroos,  the  cecum  is 

Table  13. 

Showing  the  Incidence  of  Gastroenteric  Disease  in  the  Two  Forms  of  Marsupial 

Intestinal  Tracts. 


Group  A 
Carnivorous  Stomach  and 
Intestines  103  Specimens 

Per  cent. 


Group   B 

Herbivorous  Stomach  and 

Intestines  73  Specimens 

Per  cent. 


Acute  gastritis 

Chronic  gastritis. . .  . 

Acute  enteritis 

Colitis  incl.  typhlitis 


Bacteria 
0. 


9.6 


2.9 
1.9 


Verminous 
13.6 


15. 

Bacteria  Verminous 

5.4  0. 

1.5 

4.3 


long    and    capacious    and    the    colon    relatively    long 
and  roomy. 

Table  13  contains  an  analysis  of  the  forms  of  gastro- 
enteritis as  they  were  described  in  the  two  groups  of 
tracts,  that  resembling  the  carnivorous,  that  similar  to 
the  herbivorous  intestinal  construction,  and  according  to 
the  factor  believed  to  be  responsible.  In  the  first  group 
gastritis  of  an  acute  nature  occurred  in  9.6  per  cent,  of 
the  103  specimens.  They  were  chiefly  catarrhal  in  charac- 
ter and  seem  for  the  most  part  secondary  to  verminous 
infestation ;  at  least  six  of  the  ten  cases  were  associated 
T\dth  parasites.  The  process  microscopically  is  catarrhal 
and  deeply  infiltrative.  Group  B  has  a  high  incidence  of 
gastritis  and  here  the  evidence  of  bacteria  or  toxins  is 
quite  plain.  Several  of  the  cases  were  in  animals  showing 
also  Kangaroo  disease  of  the  jaw  Avith  pneumonia  or 
septicemia;  the  gastric  lesion  of  streptothricosis  will  be 
described  under  that  heading.    The  character  of  gastritis 


THE  ALIMENTARY  TRACT  201 

without  jaAV  disease  is  somewhat  different  from  that  with 
it.  Pathologically  the  process  is  a  congestive  and  super- 
ficially necrotizing  affair,  forming  upon  the  tips  of  the 
folds,  small  gray  erosions  or  flat  shallow  irregular 
ulcers,  which  upon  histological  study  consist  of  loss  of 
tissue  of  the  mucosa  and  some  deep  congestion  with  round 
cell  groups  but  no  reaction  deep  in  mucosa  or  submucosa. 
True  catarrhal  inflammation  has  occurred,  but  not  like  in 
the  opossums. 

Chronic  gastritis  in  the  simple  stomachs  is  almost 
exclusively  in  opossums  harboring  Physaloptera  turgida, 
a  worm  which  fastens  itself  more  or  less  firmly  in  the 
mucosa  and  probably,  with  the  assistance  of  bacteria, 
causes  sufficient  irritation  to  produce  a  hypertrophic 
change  in  the  deeper  layers  and  a  destruction  of 
the  glands  where  it  holds  and  a  distortion  of  those  nearby. 
One  is  reminded  that  Fibiger  found  spiroptera  to  be 
responsible  for  adenocarcinoma  in  rats ;  no  tumor  forma- 
tion has  been  found  in  these  animals,  although  one 
opossum  with  such  a  stomach  had  an  adenocarcinoma 
mammae.  Small  hemorrhagic  spots  may  occur  in  the 
deeper  layers,  possibly  where  the  worms  have  bitten. 
The  rugae  are  irregular  or  interrupted  by  knobs 
and  papillae. 

Group  B,  stomachs  showing  chronic  change,  were  all 
kangaroos.  The  three  cases  resembled  the  infiltrating 
necrotizing  lesions  as  discussed  under  ulcers  (page  175). 
The  process  showed  an  infiltration  of  the  subsurface  tis- 
sues with  a  gray  slough  over  the  densest  part.  The 
mucosa  as  a  whole  was  irregularly  rugous  and  spotted 
with  red  gray  areas. 

Altogether  one  gets  the  impression  that  in  the  simpler 
stomach,  reactive  inflammation  is  most  prominent,  while 
in  the  colonoid  stomach  degeneration  is  greater 
than  reaction. 

Intestinal  lesions  in  marsupials  are  not  common  and 
not  peculiar  except  in  that  they  carry  out  the  pathological 

14 


202   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

reaction  as  seen  in  the  stomach.  The  carnivorous  intes- 
tine shows  frank  catarrhal  changes,  the  herbivorous 
presents  congestions  and  superficial  necroses.  These  two 
groups  then  follow  the  descriptions  as  already  given  for 
carnivores  and  ruminants. 

The  colon  presented  in  the  first  group,  simple 
catarrhal  or  follicular  change.  In  the  second  division 
the  lesions  resembled  those  in  the  stomach;  they  were 
only  once  of  the  three  instances  associated  with 
streptothricosis. 


The  class  Aves  has  been  treated  in  the  chart  precisely 
as  have  the  mammals,  but  it  is  not  possible  to  make  the 
accurate  diagnoses  or  to  separate  groups  of  lesions  ac- 
cording to  anatomy  as  was  done  for  the  latter  class.  Upon 
the  average  there  is  more  gastroenteritis  among  birds 
than  manamals,  but  when  looking  for  an  explanation  of 
this,  it  was  unfortunately  necessary  to  enter  in  the  ''unde- 
cided" column  of  Table  11  a  very  great  number  of  cases. 
This  column  would  be  greater  relatively  were  it  not  for  the 
parasites  in  parrots  and  gallinaceous  birds,  the  entries  for 
which  are  high.  Besides  this  fact  and  that  the  owl  stands 
at  the  top,  the  pigeons  at  the  bottom  of  the  list  numeri- 
cally, no  further  general  statements  are  allowable.  There 
are  several  orders  with  high  values  among  those  of  which 
insufficient  autopsies  are  at  hand  to  cast  percentages. 

The  avian  alimentary  tract  seems  to  have  developed 
according  to  the  character  of  food  the  different  varieties 
of  birds  consume  if  one  may  judge  by  the  construction  of 
the  bill,  the  gastric  musculature  and  the  intestinal  waU. 
Zoological  classification  has  only  secondarily  considered 
this  point,  it  being  made  incidental  to  the  habits  and 
habitats  of  birds.  For  this  reason  the  orders  as  outlined 
on  pages  44-46,  placed  as  they  are  in  historical  evolution- 
ary position,  represent  with  few  exceptions  groups  which 
have  differing  diets  and,  by  the  same  token,  differing  gas- 


THE  ALIMENTARY  TRACT  203 

trointestinal  tracts.  Classifications  based  upon  habits  of 
life  (Raptores,  Cantores,  Natores),  prove  likewise  too 
broad  or  too  heterogeneous,  while  systems  making 
character  of  food  the  chief  criterion  though  apparently 
correct  in  reasoning  and  helpful  in  certain  orders, 
(Accipitres,  Galli)  are  found  to  present  copious  excep- 
tions ;  moreover  we  are  imperfectly  informed  of  the  exact 
diet  that  many  families  require  or  resort  to  in  absence  of 
their  preferred  food.  I  shall  therefore  discuss  the  chief 
diseases  and  distributions  according  to  our  classification, 
preceding  the  discussion  by  a  brief  resume  of  the  ana- 
tomical peculiarities  of  the  avian  alimentary  tube. 

The  first  digestive  burden  falls  upon  the  proventricle 
where  the  principal  juices  are  secreted  while  the  muscu- 
lar stomach  or  gizzard  assumes  the  duty  of  gastric 
mastication.  The  lateral  muscular  bellies  of  its  heavy 
wall  grind  the  food  and  mix  well  the  gastric  juices.  Its 
mucosa  probably  supplies  only  lubricant.  In  birds  whose 
food  is  hard,  com  and  the  Hke,  this  grinder  is  supplied 
with  a  dry  horny  internal  layer,  while  a  thick,  moist,  soft, 
epithelial  surface  is  sufficient  for  carnivorous  birds.  All 
kinds  of  gradations  exist  between  these  extremes.  The 
mucosa  of  the  proventricle  is  always  soft,  but  quite  deep 
to  permit  the  placement  of  compound  tubular  glands.;, 

The  relation  of  size  of  these  two  parts  is  subject  to 
many  variations.  (1)  The  proventricle  is  larger  propor- 
tionately in  meat  eating,  fish  eating  and  fruit  eating 
birds,  the  gizzard  having  the  greater  size  in  granivora 
and  insectivora.  In  certain  birds  the  mucosa  of  the  two 
is  separated  by  a  very  soft  thin  zone,  an  important  fact  in 
Psittaci  since  at  this  place  spiroptera  seem  to  penetrate 
to  the  glandular  layer  of  both  organs. 

The  duodenum  begins  in  practically  all  birds,  from  a 
spherical  cavity  at  the  pyloric  end  of  the  gizzard,  to  be 
accredited  anatomically  to  both  sections.    It  passes  down- 

(1)  See  Magnan,  Compt.  Rendus  d.  I'  Acad,  de  Science,  1910  and  1911, 
Vol.  150,  151,  152. 


204  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

ward,  forms  a  long  loop  enclosing  the  pancreas,  its  distal 
end  lying  under  the  liver  and  near  the  gall-bladder.  Near 
its  end  it  receives  the  major  bile  and  pancreatic  ducts; 
smaller  ducts  from  the  liver  and  pancreas  may  enter  near 
the  pylorus  or  elsewhere  along  the  loop.  The  small 
intestine  is  usually  simple  in  its  coils,  but  in  the  birds 
that  eat  grain,  grass  and  greens,  may  be  long  and  compli- 
cated. So  too  the  colon,  usually  a  very  short  segment, 
may  be  increased  in  the  just  mentioned  group  while  the 
ceca  are  only  of  any  considerable  length  in  herbivorous 
birds.  The  length  of  the  ceca  is,  according  to  Owen, 
related  to  the  availability  of  food  and  the  need  the  bird 
may  have  for  exhausting  the  nutritive  value  of  it.  In 
carnivorous  birds  as  in  similar  manomals,  the  whole  gut, 
but  especially  the  hind-gut,  is  very  short  and  the  ceca 
small  or  absent.  But  so  they  are  in  picarian  birds  which 
are  chiefly  herbivorous,  but  may  eat  meat. 

Gasteitis. 

The  double-muscle  stomach,  that  with  the  two  lateral 
plates  and  tough  epidermal  internal  coating,  is  seldom 
the  seat  of  disease.  An  excess  of  greens  in  the  diet  some- 
times seems  to  soften  or  macerate  the  hning,  while  an 
excess  of  pebbles  may  cause  erosions.  Upon  severe  irri- 
tation this  internal  layer  assumes  the  appearance  of 
tanned  leather  and  may  crack.  The  proventricle  of  such 
a  gizzard  seems  rather  resistant  to  disease,  particularly 
one  would  say,  to  infective  processes,  for  catarrhal  or 
ulcerative  inflammation  is  uncommon.  The  saccular 
stomach  with  uniforai  muscular  walls  continuous  with 
those  of  the  proventricle,  such  as  is  seen  in  raptatory 
birds  and  parrots,  offers  a  somewhat  different  picture. 
The  internal  membranes  of  these  organs  are  definitely 
softer,  seeming  to  swell  with  great  ease,  and  the  glands 
themselves  are  smaller  both  at  the  fundus  and  outlet,  a 
construction  which  may  favor  their  closure  by  swelling 
from  simple  congestion  and  edema.    Catarrhal  and  ulcer- 


THE  ALIMENTARY  TRACT  205 

ative   processes   are   definitely  more   common   in   sucli 
organs  than  in  the  first  type  or  true  gizzard. 

Enteritis. 

Enteritis  of  all  orders  is  most  outspoken  in  the  duo- 
denal loop,  but  a  determination  of  the  lesion  must  be 
made  with  some  care.  All  the  signs  of  intestinal  inflamma- 
tion— congestion,  swelhng  and  opacity,  excess  mucus  or 
mucopurulent  covering — must  be  present  to  justify  a 
gross  diagnosis  of  enteritis  and  even  when  these  exist  one 
fails  at  times  to  confirm  the  finding  by  microscopical  sec- 
tion. These  changes  may  be  simulated  by  digestive 
activity  so  that  it  is  but  reasonable  to  demand  them  all  in 
a  clear  cut  fashion  before  applying  the  term  enteritis. 
However,  it  is  well  known  that  cases  in  both  human  and 
veterinary  medicine  giving  a  satisfactory  clinical  picture 
of  this  disease  may  fail  to  show  to  the  naked  eye  and 
under  magnification  the  changes  expected. 

In  the  hmnan  being,  the  carnivore  and  the  ruminant, 
the  ileum  presents  the  most  definite  picture  of  enteritis. 
In  the  bird,  the  duodenum  shows  the  prominent  lesions, 
and  with  the  exception  of  specific  diseases  like  blackhead, 
is  always  involved  when  the  smaller  coil  of  small  intestine 
is  affected.  This  is  true  whether  the  enteritis  be  non- 
specific or  be  associated  with  cholera  of  chickens,  ducks 
or  parrots. 

The  colon  presents  peculiar  lesions  in  but  few  birds. 
Ulcerative  and  necrotizing  processes  have  been  encoun- 
tered in  three  orders,  Psittaci,  Anseres  and  Herodiones, 
suggestively  like  the  specific  forms  seen  in  the  ceca. 
Microscopy  and  one  unsuccessful  culture  failed  to  reveal 
a  mould  or  protozoon.  The  condition  appears  as  a  gray 
white  plaque  in  the  cloacal  wall  or  it  maj^  spread  up  to  the 
colon  and  around  the  urethral  orifices.  At  times  it  is 
superficial  upon  the  mucosa  and  may  be  covered  by 
a  pseudomembrane. 


206   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

The  chief  interest  in  the  ceca  centres  about  entero- 
hepatitis  either  of  heterakis  and  amoebic  origin  or  that 
supposed  to  be  due  to  coccidia  or  Bac.  scoticus^  This 
specific  form  has  been  encountered  only  in  Galli  (three  of 
the  four  families).  It  has  been  so  well  described  by 
Hadley,  Smith,  Morse  and  Cushman  that  it  is  unneces- 
sary to  discuss  it  since  we  have  nothing  to  add  to  its 
pathogenesis  or  pathology.  At  a  later  time  some  atten- 
tion will  be  given  to  our  experience  with  Quail  disease. 
(What  is  more  interesting  from  a  comparative  stand- 
point, besides  having  a  bearing  upon  blackhead,  is 
the  discovery  of  heterakis  in  the  ceca,  and  hemorrhage 
and  fatty  change  in  the  liver  without  amoebas  or  coccidia 
in  either  place  (unfortunately  no  bacteriology  was  done), 
in  a  Sebastopol  Goose  {Anser  domesticus),  a  bird  which 
has  ceca  not  unlike  the  gallinaceous  varieties.  This  is  a 
single  observation  and  must  be  treated  expectantly. 

Microscopically  the  avian  digestive  tract  in  its  various 
inflammatory  states  presents  a  few  noteworthy  features. 
The  primary  reaction,  sometimes  the  only  one,  to  irrita- 
tion is  injection  of  the  vessels  in  the  villi  or  deeper 
mucosa.  To  this,  however,  is  nearly  always  added  a 
granularity  of  the  epithelium,  without  much  evident  mucus 
(goblet  cell)  formation.  When  the  epithelial  degenera- 
tion is  marked  there  appears  a  round  cell  increase  in 
the  deep  mucosa  shortly  followed  by  a  similar  infiltration 
into  the  villi.  True  catarrhal  enteritis  as  described  for 
the  cats  is  not  as  common  as  some  combination  of  the 
changes  just  detailed,  but  when  it  occurs  is  best  developed 
in  the  carnivorous  avian  tract.  The  most  striking  cellu- 
lar finding  is  the  round  cell  of  the  infiltrate.  It  is  of  the 
middle  lymphoid  size  with  clear  protoplasm,  or,  when  late 
in  the  disease,  may  be  small  and  so-called  adult.  Poly- 
nuclears,  unless  eosinophilic,  and  endothelioid  cells 
are  rare. 

The  foregoing  are  general  remarks  concerning  the 
pathology  of  the  avian  tract,  and  we  are  now  ready  to 


THE  ALIMENTARY  TRACT  207 

discuss  briefly  the  reactions  of  the  orders.  I  shall,  how- 
ever, omit  mention  of  those  in  the  right  half  of  Table  11. 

Passerine  birds  are  represented  better  than  any  other 
order  because  they  are  more  numerous  in  nature  and 
therefore  exhibited  more  generously  in  collections.  This 
order  is  not  especially  vulnerable  as  to  its  alimentary 
tract,  but  this  tube  is  often  affected  by  tiny  stones,  bits 
of  rust  from  cages  and  by  acute  general  non-specific  infec- 
tions to  which  these  birds  are  quite  susceptible.  Upon 
many  occasions  intense  congestion  with  and  without  tiny 
hemorrhages  in  the  duodenum  are  all  that  can  be  found  in 
the  little  birds  and  the  diagnosis  of  enteritis  is  put  down. 
Only  about  one-tenth  of  these  birds  have  shown  more  or 
less  definite  catarrhal  changes.  Many  birds  have  heavy 
infestation  with  worms  which  may  activate  bacteria. 

The  Striges  and  the  Accipitres  will  be  discussed 
together  because  of  the  similarity  of  their  tracts  and  their 
diets.  The  reaction  of  their  gastric  complex  has  already 
been  mentioned  and  what  was  written  there  can  be 
extended  to  the  intestine.  The  type  of  lesion  is  catarrhal 
and  seems  to  be  "meat  bred"  although  this  cannot  be 
proven.  They  never  have  given  positive  heart's  blood 
cultures  so  that  the  disease  seemed  not  to  be  septicemic. 
Liver  and  spleen  have  harbored  colon  and  paracolon 
bacilli.  It  will  be  noticed  that  they  have  the  highest 
incidence  of  gastric  disease. 

Psittaci,  birds  with  a  tract  similar  to  the  last  two  but 
with  a  captive  diet  of  seeds,  fruit  and  vegetables 
although  they  may  eat  small  animals  and  insects  in  the 
wild,  present  figures  under  bacteria  and  parasites  which 
explain  the  involvement  of  the  alimentary  organs.  We 
have  had  two  acute  outbreaks  of  what  seemed  to  be  fowl 
cholera,  judging  by  the  pathology  and  the  isolation  of 
Bad.  gallinarum  and  we  have  frequent  deaths  with  the 
same  gross  anatomy  from  which  bacterial  isolation  has 
not  been  tried  or  was  unsuccessful.  At  all  events  the 
enteritis  of  parrots  is  often  a  definitely  infectious  affair. 


208   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

We  have  also  isolated  Bad.  psittacosis  on  two  occasions. 
Separation  of  the  birds  and  cleansing  of  the  exhibition 
spaces  seemed  to  stop  the  disease.  The  lesions  are 
hemorrhagic  and  superficially  necrotizing  in  this  group 
whether  or  not  they  are  septicemic.  Some  have  also 
sho^vn  a  follicular  appearance  both  grossly  and  minutely, 
one  of  which  cases  was  associated  with  Bad.  psittacosis. 

The  effect  of  animal  parasites  is  chiefly  exerted,  for 
this  group,  in  the  proventricle  where  the  Spiroptera 
incerta  occupies  the  lumen  and  penetrates  the  mucosa  or 
burrows  under  the  inner  lining  of  the  gizzard.  Enteritis 
is  not  especially  associated  with  this  infestation,  death 
resulting  more  from  inanition  than  infection;  some  few 
cases  have  had  enteritis,  others  pulmonary  disease. 

Columbai  are  not  susceptible  to  disease  in  the  parts 
under  consideration.  This  order  seems  to  have  some 
tissue  resistance,  for  their  lesions  are  quite  frankly 
catarrhal,  more  so  than  in  most  grain-eating  birds. 

The  figures  set  against  the  Galli  are  swollen  by  the 
number  of  cases  of  enterohepatitis  of  one  sort  or 
another.  Extracting  these  from  the  total  leaves  the  order 
among  the  lowest.  Their  lesions  are  congestive  and 
hemorrhagic,  although  they  may  show  catarrhal  cases. 
They  seem  to  be  able  to  summon  mucus  more  readily  than 
many  other  birds. 

Anserine  birds,  though  not  very  high  in  figures,  pre- 
sent three  conditions  worthy  of  note.  In  the  first  place, 
acute  simple  gastritis  occurs  often,  sometimes  associated 
with  foul  green  food,  sometimes  without  any  apparent 
cause.  From  the  number  of  times  that  foreign  bodies  are 
present  it  seems  probable  that  they  contribute  in  some 
Avay.  Excessive  stones  and  sand,  bits  of  glass,  collar 
buttons  and  the  like  are  sometimes  found.  Then  the  form 
of  acute  enteritis  has  always  a  hemorrhagic  tendency,  at 
least  in  the  submucosa,  while  the  mucosa  may  be  swollen, 
opaque  and  covered  with  mucus.  Upon  histological  study 
these  intestines  show  intense  swelling  by  cellular  infil- 


THE  ALIMENTARY  TRACT  209 

trate  and  disappearance  of  the  tips  of  the  villi.  The  third 
observation  concerns  what  is  apparently  a  subacute  or 
chronic  process  although  this  is  not  supported  by  micros- 
copy. Certain  birds  will  have  a  cast  of  mucus  and 
epithelial  detritus  rather  closely  adherent  to  the  wall. 
Under  the  microscope  there  may  be  slight  evidence  of 
chronic  inflammation  or  there  may  be  little  amiss.  These 
birds  have  usually  been  large  ones,  and  several  have  come 
from  the  separate  goose  pens,  not  from  the  open  lake 
where  many  birds  are  kept. 

The  struthious  birds  deserve  a  word.  They  have  had 
a  great  deal  of  enteritis  and  mostly  of  infectious  nature. 
Two  instances  have  arisen  from  bird  diphtheria,  one  from 
cholera  and  six  from  what  later  seemed  to  have  been 
anthrax  but  was  not  diagnosed  at  the  time.  The  character 
of  the  lesions  in  the  struthious  intestine  tends  to  be 
hypertrophic  and  superficially  erosive  if  not  ulcerative. 
The  changes  are  found  with  greatest  clearness  in  the 
lower  duodenum  and  small  coil. 

Constipation. 

(Having  discussed  the  inflammatory  conditions  of  the 
gastrointestinal  tract  we  now  come  to  the  more  or  less 
definitely  mechanical  abnormalities,  whether  or  not  they 
depend  upon  preexisting  inflammation,  and  the  subject  of 
constipation  will  claim  first  attention.  In  the  human 
Jbeing  this  condition  is  the  result  of  bad  habits  more  than 
any  other  one  thing  or  all  things  together,  I  think  it  will 
be  admitted.  In  the  lower  animals  perhaps  no  such  thing 
as  habit  of  defecation  exists  so  that  one  can  with  more 
certainty  hold  incorrect  food,  chronic  catarrhs  or  physical 
obstruction  as  responsible.  Veterinarians  look  upon 
excess  of  dry  food  and  irregularity  of  work  and  food 
periods  as  the  principal  causes  of  constipation.  These 
factors  do  not  hold  in  zoological  collections.  As  a  matter 
of  fact  constipation  is  of  minor  importance  in  this 
menagerie,  but  a  certain  few  cases  are  worthy  of  note.    It 


210  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

has  been  mentioned  in  the  diagnoses  in  only  a  little  over 
1  per  cent,  of  the  total,  and  of  these  the  records  indicate 
its  importance  only  ten  times  (.2  per  cent.) ;  a  few  notes 
of  these  cases  are  appended.  The  first  place  of  incidence 
is  taken  by  marsupials  (six  kangaroos  and  one  opossum), 
the  second  by  ungulates  (largely  ruminants)  and  the 
third  by  Primates.  It  will  be  noted  that  with  exception 
of  the  opossum,  herbivorous  mammals  occupy  the  first 
places  of  incidence,  carnivores  falling  well  behind  the 
orders  named.  This  condition  is  quite  infrequent  in 
birds  and  is  usually  associated  with  the  presence  of  seeds 
or  parasites  or  with  impaction  in  the  ceca. 

Primates,  almost  exclusively  feeders  upon  carbo- 
hydrate and  soft  protein  food,  have  shown  as  causes  of 
constipation  two  outstanding  conditions.  A  low  grade 
of  colonic  catarrh  with  excessive  pouchings  of  this  tube 
has  had  constipation  associated  with  it  three  times.  One 
of  these  cases  had  small  coproliths  in  the  diverticula,  one 
other  a  fecal  concrement  in  the  cecum.  Another  group  of 
these  cases  with  evidence  of  delayed  passage  of  feces 
shows  chronic  peritonitis  with  adhesions,  one  of  which 
seems  certainly  due  to  filaria  in  the  peritoneal  sac. 

The  seat  of  constipation  in  monkeys  is  practically 
always  the  colon.  The  carnivores  while  occasionally 
showing  hard  fecal  masses  packed  into  the  colon,  more 
often  exhibit  a  constipation  in  the  ileum.  One  case  pre- 
sented a  nearly  empty  colon  with  a  long  scybalum  just 
above  the  cecal  valve.  There  is  no  peculiar  associated 
pathology  in  the  notes  at  my  command. 

Ungulata,  showing  next  to  the  highest  incidence,  has 
its  stoppage  chiefly  in  the  colon,  but  the  lowest  stretch  of 
the  ileum  may  contain  balls  of  feces.  In  nearly  every  case 
one  finds  some  grade  of  colonic  catarrh.  In  two  instances, 
there  being  a  proctitis,  it  seemed  as  if  the  animal  volun- 
tarily restrained  from  defecating  because  of  pain.  The 
caput  coli  is  the  seat  of  stoppage  in  the  Rodentia. 


THE  ALIMENTARY  TRACT  211 

Marsupials  give  such  a  high  relative  incidence  that 
especial  search  of  their  records  was  made,  Avithout,  how- 
ever, very  definite  result.  In  three  of  the  seven  cases  an 
acute  general  infection  existed,  in  one  an  acute  peritonitis 
which  seemed  to  emanate  from  a  small  ulcer  in  the  ileum, 
in  one  an  injury  to  the  anal  region  was  found  while  in  the 
remaining  two  the  notes  would  suggest  that  the  lower 
intestine  was  atonic,  judging  by  its  distention,  trans- 
lucency  and  pallor.  In  five  the  stoppage  took  place 
in  the  large  bowel  alone,  in  the  others  both  divisions 
being  affected. 

It  is  often  difficult  to  establish  a  diagnosis  of  consti- 
pation in  birds  because  many  varieties  form  a  long  rather 
dry  mass  in  the  lower  small  intestine,  to  be  moistened  in 
the  cloaca  for  discharge.  Still  again  the  groups  with 
capacious  ceca  are  apt  to  have  them  filled  normally  with 
firm  casts.  Diagnoses  of  fecal  inspissation  and  stoppage 
in  the  smaller  tube  have  been  made  seldom,  but  one  must 
consider  also  the  obstruction  offered  by  excessive  urate 
collections  either  in  the  cloaca  or  lower  ileum  which  will 
amount  to  a  constipation  if  the  cloaca  be  over-dilated 
and  dried  urates  mixed  with  dirt  or  feathers  cover  the 
anal  opening. 

The  causes  of  this  condition  in  birds  are  usually 
mechanical,  inflammation  being  found  in  a  small  minority 
of  cases.  In  the  small  passerine  birds,  seeds,  sand,  or 
parasites  form  the  commonest  findings.  This  is  also  true 
of  parrots,  while  excessive  urate  collections  are  noted  for 
both  these  groups.  The  gallinaceous  birds  present  two 
reasons  for  fecal  stoppage — disease  of  the  ceca  (see 
pages  205-6)  and  cloacitis  probably  secondary  to  anal 
closure  by  excessive  urate  collection.  Uratic  stones, 
varying  from  one  to  five  millimetres  in  diameter,  have 
been  found  in  the  cloaca  in  several  orders.  In  only  one 
case,  a  pheasant,  did  they  cause  ulceration  and  cloacitis. 
Sand,  rust,  grains  and  the  like  are  found  frequently,  and 
sometimes  in  groups  of  birds,  indicating  that  the  speci- 


212   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

mens  had  not  been  put  upon  the  correct  flooring  or 
caging.    Unbroken  seeds  may  obstruct  the  lumen.y 

Mechanical,  Obstruction. 

Although  the  following  is  not  constipation  it  is  well  to 
cite  at  this  place  an  experience  which  amounted  to 
mechanical  intestinal  obstruction.  A  number  of  finches 
were  subjected  to  postmortem  and  found  to  have  whole 
white  millet  seeds  in  their  intestines,  this  being  the  only 
discoverable  cause  of  death.  Investigation  revealed  that 
during  the  night  mice  ate  the  canary  seed  in  the  pans, 
leaving  only  the  millet,  w^hich  the  hungiy  birds  consumed 
w^hole.  Small  birds  can  take  a  few  millet  and  crack  them 
when  eating  leisurely,  but  apparently  not  when  hungry. 
When  the  food  was  removed  at  night  the  trouble  ceased. 

Obstruction  by  sand  is  well  illustrated  by  a  peculiar 
form  of  pica,  in  a  goose,  which  is  worth  citing,  and  calls  to 
mind  the  sand  disease  of  horses : 

Canada  Goose  6    {Branta  canadensis  canadensis). 

Diagnosis. — Masses  of  sand  in  entire  intestinal  tract.  The  general 
condition  externally  and  internally  is  good.  The  crop  is  distended  like 
a  sausage,  quite  firm  and  the  overfilling  is  obviously  due  to  sand  in 
which  veiy  few  stones,  Avhieh  could  be  called  pebbles,  are  found.  This 
mass  continues  into  the  esophagus  making  the  whole  tract  impassable 
for  food.  The  mucosa  is  a  little  pink  and  dirt-stained  in  places  but  is 
not  visibly  inflamed.  The  gizzard  is  contracted  over  a  mass  of  sand 
but  no  food.  Sand  in  more  or  less  definitely  packed  condition  is  found 
all  along  the  gut  tract,  in  one  place  in  the  small  coil  it  being  quite  as 
tight  as  in  the  crop  and  no  lumen  remaining.  Sand  and  bits  of  shale 
are  found  in  ceca.  The  organs  are  apparently  healthy,  slightly  pale 
perhaps,  but  certainly  not  distinctly  anemic.  No  infection  exists.  The 
aorta,  just  above  renals,  has  a  15  ram.  x  2  mm.  pale  opacity  of  same 
consistency  as  the  rest  of  the  vessel,  just  perceptibly  higher  than  sur- 
rounding surface. 

*'Sand  disease"  has  occurred  in  a  Persian  Wild  Ass 
(Equus  onager)  causing  in  this  case  ulceration,  perfora- 
tion and  peritonitis,  a  Common  deer  {Mazama  virginiana) 
and  a  Chapman's  zebra  {Equus  hurchelU  cliapmani). 
The  collection  of  sand  is  always  greatest  in  the  caput  coli, 
but  may  coat  the  large  bowel  to  the  anus. 


THE  ALIMENTARY  TRACT  213 

Larger  and  more  definitely  obstructive  physical 
objects  are  found  in  both  mammals  and  birds.  We  have 
on  record  a  lion  (Felis  leo)  and  a  tiger  {Felis  tigris), 
which  swallowed  pieces  of  bone  large  enough  to  be  stuck 
in  the  small  intestine  and  completely  occlude  it.  Smaller 
objects  like  buttons  have  been  found  even  in  the  passerine 
tract.  Worm  masses  may  occupy  such  a  large  part  of  the 
lumen  of  the  tube  as  to  constitute  a  physical  obstruction. 
This  is  definitely  less  important  in  mammals  than  in 
birds,  especially  in  the  passerine  order  of  the  latter  class. 

Dilatation  of  the  intestine  aside  from  that  occurring 
in  connection  with  fermentation,  constipation  or  ileus,  in 
other  words  chronic  atonic  dilatation,  has  not  been 
encountered.  Acute  dilatation  has  been  found  in  several 
orders  under  the  picture  known  for  domesticated  animals. 
Its  pathology  and  incidence  have  already  been  discussed. 

Ileus. 

(^  Ileus  or  acute  intestinal  obstruction  may  be  divided 
for  our  purposes  into  intussusception,  volvulus,  strangu- 
lation and  paralysis  from  interruption  of  mesenteric 
circulation.  Examples  of  all  these  varieties  have  been 
encountered  and  illustrative  cases  will  be  cited.  In  so  far 
as  incidence  is  concerned,  the  Ungulata  and  Carnivora 
greatly  outnumber  all  other  orders,  showing  seven  cases 
each ;  the  sum  total  in  all  other  orders  is  but  eight.  Upon 
re-reading  some  of  the  protocols  I  have,  however, 
excluded  three  invaginations  in  the  carnivores,  one  each 
in  the  ungulates  and  rodents,  as  probably  being  post- 
mortem or  shortly  antemortem  occurrences ;  two  had  very 
early  peritonitis  but  other  things,  sufficient  to  account  for 
death,  were  present.  These  deductions  bring  the  total 
cases  of  ileus  in  mammals  to  seventeen.  Five  cases  in 
birds  will  be  discussed  briefly. 

Primates  present  one  case  of  volvulus,  one  of  intus- 
susception and  one  of  internal  strangulation.  The  first 
displayed  the  entrance  of  four  inches  of  ileum  into  the 


214   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

colon  with  such  swelling  of  the  waU  as  to  prevent  reduc- 
tion. The  exciting  cause  seemed  to  be  an  enteritis,  the 
cause  of  death  a  peritonitis.  A  white-collared  mangabey 
{Cercocebus  collaris)  was  the  victim  of  volvulus  probably 
favored  by  an  anomalous  position  of  the  transverse  and 
descending  colon  which  lay  to  the  right,  the  latter 
traversing  the  abdomen  obliquely  from  right  to  left  to 
reach  the  pelvis.  The  volvulus  occurred  in  the  ileum  just 
above  the  cecum,  the  tmsted  part  being  found  adherent 
by  the  peritonitis.  The  third  case  is  a  strangulation  due 
to  peritonitis  from  filaria  and  adliesions  between  stomach 
and  colon  from  a  colitis  and  pericolitis  due  to  cestodes, 
one  of  which  was  found  deeply  implanted  in  the 
colonic  wall. 

Two  cases  of  intussusception  are  noted  (after  deduc- 
tions above)  for  the  Carnivora.  They  both  occurred  in 
the  ileum,  one  restricted  thereto,  the  other  extending  into 
the  colon.  In  both  a  vague  history  of  being  ''  off  their 
feed  "  or  giving  evidence  of  intestinal  trouble  could  be 
obtained  from  the  keeper.  The  three  excluded  cases  had 
invaginations  in  the  middle  and  lower  small  intestines 
but  not  at  the  cecal  valve.  Volvulus  did  not  occur  in 
the  Carnivora. 

Aparadoxure  {Paradoxurus  hermophroditus)  died  as 
the  result  of  a  strangulation  of  a  six-inch  knuckle  of  gut 
which  had  passed  through  a  hole  in  the  omentum.  The 
animal  had  not  been  eating  well  for  a  month  but  gave  no 
signs  by  which  this  ileus  could  have  been  diagnosed.  Per- 
haps it  had  existed  for  sometime  but  only  shortly  before 
death  had  swollen  sufficiently  to  cause  obstruction. 

Having  excluded  a  doubtful  invagination  in  a  small 
rodent  there  remains  an  interesting  though  somewhat 
obscure  case  in  a  porcupine  {Erethizon  dorsatus  dor- 
satus).  This  animal  suffered  with  an  acute  hemorrhagic 
and  catarrhal  enteritis  while  the  colon  seemed  free  of 
change  until  the  rectum  was  reached.  Here  was  a  stretch 
of  a  foot  with  the  purple,   lusterless  but  translucent 


THE  ALIMENTARY  TRACT  215 

appearance  of  a  strangulated  intestine  although  no  invo- 
lution or  twisting  remained.  This  was  looked  upon  as 
a  volvulus  which  had  untwisted  a  few  hours  before  death. 

Intussusception  was  seen  only  once  in  the  Ungulata, 
a  tapir  (Tapirus  terrestris)  with  chronic  enteritis.  Here 
the  ileum  had  passed  into  the  colon  for  a  distance  of  nine 
inches,  it  being  much  swollen  and  congested  but  not 
gangrenous.  Its  condition  warranted  the  idea  that  the 
process  was  antemortem  but  a  peritonitis  had  not  arisen, 
death  having  occurred  from  the  slight  extra  shock  in  an 
animal  suffering  with  chronic  malnutrition.  Volvulus 
was  encountered  three  times,  two  deer  and  a  zebra.  The 
last  was  the  animal  already  described  that  carried  such  a 
heavy  load  of  sand  in  the  gut  tract,  a  factor  in  the  produc- 
tion of  the  twist  probably  although  this  might  have  been 
aided  by  a  fibromyoma  of  the  colonic  wall.  The  location 
of  the  volvulus  in  this  order  was  twice  in  the  dilated 
descending  colon,  the  third  in  the  jejunal  area.  This  last 
was  a  twist  which  resembled  an  internal  strangulation 
because  of  the  intricate  knot-like  windings  of  the 
smaU  bowel. 

The  marsupials  present  two  interesting  cases.  A  rock 
kangaroo  {Petrogale  pencillata)  had  chronic  gastric 
ulcerations  with  local  peritoneal  adhesions  which  appar- 
ently obstructed  nearby  coils  of  intestine  so  that  they 
became  inflated  and  twisted  over.  An  opossum  had  a  vol- 
vulus of  the  stomach  which  performed  one  and  a  half 
turns  from  left  to  right ;  its  protocol  follows. 

Common  Opossum  6  (Didelphys  virginiana).  Ileus.  One  and  one- 
half  complete  volvulus  turns  of  stomach  on  duodenum.  General  condi- 
tion fairly  good.  Abdomen  quite  prominent,  a  condition  found  to  be  due 
to  great  dilatation  of  the  stomach  which  occupied  the  whole  anterior 
part  of  the  abdominal  cavity.  The  organ  is  blue  and  the  vessels  stand 
out.  Postmortem  changes  are  occurring  everywhere  favored  by  the 
obstruction  to  the  circulation.  The  dilated  stomach  has  undergone  a 
volvulus  upon  the  third  part  of  the  duodenum  making  one  and  a  half 
turns.  The  spleen  lies  upon  the  right  side  well  below  the  liver;  it  is 
swollen,  soft  and  deep  purple.  The  duodenum  in  its  upper  half  takes 
part  in  the  dilatation  and  beginning  gangrene.     The  pedicle  of  the 


216  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

twist  is  made  of  the  duodenum,  esophagus,  edge  of  the  mesentery  and 
the  middle  part  of  the  pancreas;  the  end  of  the  tail  of  the  last  is  gan- 
grenous. There  is  no  apparent  obstruction  lower  down  to  explain  the 
twist  of  the  stomach. 

Among  the  Aves  the  following  cases  only  are  worthy 
of  report.  A  parrot  {Melopsittacus  undulatus)  was  found 
to  have  a  tightly  packed  mass  of  worms  in  the  end  of  the 
duodenum  above  wliich  the  bowel  was  distended,  enlon- 
gated,  doubled  on  itself  and  of  a  deep  red  color ;  below  this 
the  small  intestine  was  empty.  A  closely  siioilar  con- 
dition was  found  in  a  Screech  Owl  {Otus  asio  asio)  the 
obstruction  occurring  just  above  the  end  of  the  small  gut. 
A  Sparrow  Hawk  {Falco  sparverius)  had  an  invagination 
two  cm.  in  length,  a  short  distance  above  the  end  of  the 
small  intestine.  No  peritonitis  existed  but  the  presence 
of  an  acute  enteritis  helps  to  explain  the  intussusception. 

Hernia. 

Hernia  is  not  a  common  occurrence  among  the  lower 
animals  but  our  experience  is  instructive  in  two  partic- 
ulars, to  wit,  its  absence  in  the  orders  preceding  the 
Eodentia  and  the  frequency  of  the  traumatic  variety. 
There  being  no  general  remarks  to  be  made  upon  the  sub- 
ject, it  seems  well  to  give  a  summary  of  the  findings  in 
each  of  the  seven  cases. 

A  Western  Fox  Squirrel  (Sciurus  rufiventer)  showed 
a  diaphragmatic  defect  on  the  right  side,  a,  rounded  open- 
ing with  smooth  edges,  through  which  a  loop  of  intestine 
had  passed,  entering  behind  the  liver  and  reaching  into  the 
pleura  as  high  as  the  pulmonary  apex  where  it  was 
adherent ;  this  was  probably  of  long  standing.  Two  more 
loops  were  found  wedged  in  the  diaphragmatic  hole,  one 
of  which  was  gangrenous. 

An  Indian  Antelope  {Black  Buck)  {Antilope  cervi- 
capra)  presented  an  irreducible  incarcerated  but  not 
strangulated  umbilical  hernia.  The  peritoneum  was  fused 


THE  ALIMENTARY  TRACT  217 

with  the  aponeurosis  at  the  ring  but  the  gut  was  not 
adherent  at  this  point  while  it  was  attached  within  the 
sac  outside  the  muscle,  thus  forming  the  incarceration. 
Apparently  the  sac  had  dissected  between  the  muscular 
layers  for  it  could  be  followed  for  several  centimetres  in 
some  directions.  A  Hog  Deer  {Cervus  porcinus)  had 
apparently  suffered  an  injury  in  the  flank  for  at  one  point 
the  muscles  were  irregularly  cicatrized  and  a  rent  was 
present  through  which  several  loops  of  intestine  and  a 
band  of  omentum  had  escaped,  being  adherent  to  fascia. 
No  injury  to  the  skin  was  apparent. 

Another  Indian  Antelope  showed  a  clean  traumatic 
rupture  of  the  muscle  mid  peritoneum  in  the  right 
inguinal  region  ivithout  penetration  of  skin.  An  acute 
hernia  had  occurred  which  was  lightly  adherent  to  fascia 
and  an  acute  peritonitis  was  beginning.  The  bowel  was 
however  not  strangulated. 

An  aoudad  {Ovis  tragelaphus)  seems  to  have  suffered 
an  injury  hy  a  pointed  object  (horn?)  just  to  the  right  of 
the  ensiform  cartilage  for  at  this  position  there  is  a 
circular  hole,  with  smooth  healed  edges,  in  the  aponeuro- 
sis, permitting  the  emersion  of  a  peritoneal  sac  contain- 
ing omentum.  All  parts  were  adherent  but  no  acute 
inflammation  existed. 

( What  may  have  been  a  hernia  or  a  relaxation  of  the 
transversus  perinei  was  observed  in  an  Undulated  Grass 
Parrakeet  {Melopsittacus  undulatus).  A  bulge  about  the 
size  of  the  finger  end  was  seen  externally,  beside  and 
behind  the  anus.  This  proved  to  contain  several  loops  of 
bowel  and  a  mass  of  fat. 

A  lateral  abdominal  hernia  was  seen  in  a  Barbary 
Turtle  Dove  (Turtur  risorius).  It  consisted  of  a  per- 
itoneal sac  and  two  loops  of  intestine.  This  protrusion, 
while  firmly  fixed  in  its  unnatural  position,  was  in  no 
way  constricted. 

15 


218   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Rectal,  Prolapse. 

Prolapse  of  the  rectum  may  in  a  sense  be  looked  upon 
as  a  hernia  or  at  least  as  a  relaxation  of  the  anal  and 
perineal  muscles  with  protrusion  of  parts  normally  situ- 
ated intracorporeally.  Although  not  frequent  it  has  been 
incurable  in  the  animal,  as  it  frequently  is  in  man  without 
operation,  a  measure  we  have  not  adopted.  Just  what 
determines  weakness  in  the  pelvic  outlet  is  entirely 
obscure  for  indeed  we  have  seen  here  wounds  and  inflam- 
mations of  the  perineal  area  \vithout  prolapse  of  the 
rectum  and  in  none  of  the  cases  of  prolapse  did  the  pelvic 
floor  seem  injured  or  diseased.  It  is  but  speculation  to 
blame  the  annular  muscles  of  the  anus.  Tenesmus,  or  at 
least  reasons  for  this  straining  action,  have  been  sought, 
with  the  result  that  in  our  cases  lesions  of  the  egg-laying 
apparatus  in  birds  and  enteritis  in  mammals  have  stood 
out  most  prominently.  In  no  case  have  hemorrhoids  been 
encountered  nor  has  a  tumor  pendant  from  the  colonic 
mucosa,  drawn  the  bowel  toward  the  anal  opening.  It 
might  be  added  parenthetically  here  that  hemorrhoids  are 
practically  unlaiown  for  quadrupeds,  Hutyra  and  Marek 
failing  to  mention  them  independently  and  only  one  refer- 
ence being  found  in  the  Jahresbericht  fur  Veterindr 
Medizin  (Schmidt  1914-169);  this  case  is  more  like 
angioma  than  hemorrhoids.  If  tenesmus  be  active  in  the 
production  of  rectal  prolapse  then  it  would  have  to  be 
assumed  that  this  straining  effort  can  be  induced  by  enter- 
itis since  eversion  of  the  rectum  has  occurred  with  this 
disease  in  the  absence  of  colitis,  the  condition  usually 
expected  in  the  presence  of  tenesmus.  The  thirteen  cases 
have  been  seen  in  Mammalia,  8,  (Carnivora,  2,  Rodentia,  1, 
Ungulata,  3,  Edentata,  1,  Marsupialia,  1)  and  Aves,  5, 
(Passeres,  Picari.T',  Striges,  Psittaci  and  Galli  each  one). 
Three  mammals  had  enteritis,  one  had  foreign  bodies  in 
the  bowel  and  one  had  many  ascarids;  three  had  no 
demonstrable  or  suggestive  causes.     Two  of  the  five  birds 


THE  ALIMENTARY  TRACT  219 

had  enteritis  high  in  the  tract,  one  had  uratic  calculi  in 
the  cloaca,  and  three  had  trouble  in  the  egg  laying 
apparatus :  one  too  large  an  egg,  one  a  broken  egg  and 
one  a  salpingitis.; 

Diverticula. 

It  is  almost  certain  that  in  a  human  pathological 
service  of  fifty-five  hundred  autopsies,  one  or  more 
diverticula  of  the  Meckel  variety  would  be  encountered 
and  perhaps  several  of  other  kinds.  In  our  material 
only  pouchings  or  false  diverticula  of  the  colonic  wall 
are  recorded,  and  our  personnel  has  often  spoken  of 
the  absence  of  these  gross  abnormalities  of  the  alimentary 
tract.  The  two  cases,  notes  of  which  are  given,  are 
instances  of  hernial  pouchings  of  the  colonic  mucosa  and 
serosa,  a  condition  which  is  well  known  in  human  medi- 
cine. C,It  may  be  said  to  occur  in  two  varities,  one  in  which 
the  pouchings  have  heavy  walls  formed  by  a  thickened 
mucosa,  muscularis  and  peritoneum  and  on©  in  which  the 
bulgings  have  delicate  walls,  then  being  small  herniae  of 
the  inner  coats  through  rifts  in  the  outer.  Such  a  division 
is  probably  unnecessary  or  misleading  since  the  latter 
may  be  only  a  forerunner  of  the  former.  However  the 
clinical  evidence  of  the  simple  variety  is  scanty  and  may  be 
little  more  than  constipation  while  the  peritonitic  variety 
gives  a  clinical  picture  of  pain,  constipation  and  a  mass  in 
the  left  abdominal  area,  then  kno^vn  as  diverticulitis  or 
pericolitis  sinistra.  In  these  cases  the  colon  is  much  dis- 
torted by  the  irregularity  of  its  mucosa  and  by  inflamma- 
tory thickening  of  the  muscularis  and  serosa.  Diverticula 
arise  from  defects  of  the  muscular  coat,  or  second- 
arily after  inflammation  or  prolonged  constipation,  by 
weakness  of  muscle,  or  as  hernial  protrusions  around  the 
entrance  of  blood  vessels  where  the  muscle  is  thin.  Such 
sacculations  permit  feces  to  collect  and  continue  the 
inflammation,  thus  further  weakening  the  gut  and  pro- 
ducing constipation,  the  whole  vicious  cycle  being  favor- 


220   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

able  to  the  formation  of  more  sacculations;  coproliths 
may  form  in  the  diverticula.  The  two  monkeys  now 
reported  seem  to  have  varying  grades  of  the  same  con- 
dition, a  long  standing  colitis  \\ith  diverticula,  constipa- 
tion and  the  collection  of  inspissated  feces  in  the 
sacculations.    These  animals  did  not  have  hemorrhoids. 

Black  Ape  9  {Cynopithecus  niger).  Coprostasis.  Coproliths  in 
diverticulum.  Chronic  colitis.  Cor  bifida.  The  large  intestine  is  of 
the  same  calibre  as  the  small  intestine  should  be  when  not  distended. 
The  sacculations  as  seen  before  opening  the  organ  are  salient,  forming 
distinct  pouches.  In  one  or  two  cases  they  are  so  pronounced  as  to 
constitute  diverticula  7  cm.  long.  In  two  instances  the  serosa  at  the 
fundi  of  these  diverticula  is  markedly  hyperemie  and  very  thin.  In 
many  cases  the  sacculations  contain  coproliths.  The  wall  of  the  organ 
is  distinctly  thickened,  puckered,  inelastic  and  opaque.  Mucosa  is 
thrown  up  into  coarse  rugae. 

Japanese  Macaque  6  (Macacus  fuscatus).  Chronic  hypertrophic 
colitis.  False  diverticula  of  colon.  The  large  intestine  contains  a 
moderate  quantity  of  quite  constipated  feces.  The  serosa  is  smooth. 
The  wall  shows  at  several  stretches  enlargements  of  the  normal  saccu- 
lations, forming  false  diverticula.  The  wall  of  the  gut  in  these  herniae 
is  thinner  than  in  the  surrounding  parts;  no  ulcers  exist;  no  local 
peritonitis  is  present.  The  mucosa  everywhere  is  irregular  in  thickness, 
less  translucent  than  normal  and  thrown  into  irregular  rugaB;  tenacious 
mucus  covers  it.     No  ulcers. 

TUMOES. 

Only  one  tumor  was  observed  in  the  mammalian  intes- 
tinal tract  proper.  Dasyure  {Dasyuriis  macidatus) 
Adenocarcinoma  of  the  intestines.  On  postmortem  there 
was  a  pale  diffuse  thickening  of  the  coats  of  the  small  gut 
over  a  large  area;  numerous  soft,  light  yellow,  sharply 
circumscribed,  elevated  (like  secondary  tumors)  nodules 
in  the  liver  and  spleen,  and  a  pea-size  whitish  nodule 
around  a  bronchus  in  the  right  lung.  Histological  section 
of  primary  growth  not  made  but  a  cross  section  of  the 
intestine  in  the  vicinity  shows  an  adenomatous  change 
with  considerable  increase  in  the  connective  tissue.  The 
nodules  in  the  liver,  spleen  and  lung  and  the  appearances 
of  the  abdominal  IjTnph  nodes,  found  microscopically,  are 
precisely  similar.     They  consist  of  irregnilarly  arranged 


THE  ALIMENTARY  TRACT  221 

epithelial  nests  and  distorted  acini,  around  which  are 
sharply  outlined  spaces,  filled  with  the  remains  of  de- 
generated blood  or  a  granular  material.  The  metastases 
are  always  sharply  outlined. 

Aves  supply  three  papillomata  which  are  interesting 
in  that  one  occurred  in  the  proventricle,  and  two  grew  in 
the  duodenum  in  the  vicinity  of  the  upper  biliary  opening 
and  presented  within  the  lumen  soft  masses  which,  while 
not  occluding  the  passage,  offered  some  little  obstruction 
as  indicated  by  a  slight  distention  above  their  location.  In 
two,  carefully  studied,  no  indications  of  parasites  or  of 
cancer  could  be  found.  The  birds  concerned  were  an 
amazon,  an  owl,  and  a  rhea. 


SECTION  VII 
THE  ALIMENTARY  TRACT,  PART  2.  THE  LIVER 

A  consideration  of  the  liver  is  anatomically  and 
physiologically  the  next  step  in  the  discussion  of  diseases 
of  the  alimentary  tract.  While  this  organ  may  partici- 
pate in  most  of  the  pathological  states  of  the  tubal  part  of 
the  system,  it  is  comparatively  seldom  the  primary  seat 
of  change  and  when  damaged  seems  to  be  possessed  of 
great  accommodative  and  reconstructive  power.  This 
must  be  true,  and  fortunately  so,  since  we  ascribe  to  it 
the  major  detoxicating  function  of  the  body.  Neverthe- 
less it  is  noteworthy  that  the  largest  solid  organ  of 
the  animal  body  shows  a  relatively  low  percentage  of 
changes  threatening  to  life.  In  the  sense  of  Pearl's 
method  of  statistics,  it  does  not  ''  break  down  "  easily. 
In  pathology  it  is  the  custom  to  list  with  great  care  all  the 
changes,  gross  and  minute,  in  the  liver,  but  with  a  few 
exceptions  they  are  secondary  or  incidental.  They  do 
however  reflect  many  things,  especially  referable  to  diet 
and  to  chronic  infection  from  the  intestinal  drainage 
area.  It  is  in  these  directions  that  the  organ  will  be 
studied  in  the  following  pages. 

Anatomically  the  liver  is  situated  in  the  right  upper 
part  of  the  abdomen  subjacent  to  the  diaphragm  in  both 
mammals  and  birds,  being  held  in  position  by  attachment 
to  this  transverse  partition,  by  ligaments  or  folds  of 
peritoneum,  and  by  the  other  abdominal  viscera.  Its 
general  relationships  do  not  offer  great  variations  since 
in  all  animals  means  are  afforded  for  a  dual  blood  supply 
and  an  outlet  for  the  hepatic  secretion,  the  bile,  into  the 
higher  intestines.  Naturally  variations  in  the  size  of  the 
lobes  are  observed  and  there  has  been  considerable  spec- 
ulation as  to  their  independence  and  association.  From 
the  standpoint  of  comparative  pathologj%  little  can  be 

222 


THE  LIVER  223 

ascertained  to  assist  in  this  matter  unless  the  position  of 
abscesses  and  hepatitis  relative  to  cholecystitis  have  a 
bearing;  some  discussion  of  this  will  appear  later.  We 
have  not  observed  any  peculiar  pathology  of  the  lobes  of 
Spigelius  and  Riedel.  In  so  far  as  the  size  and  arrange- 
ment of  the  organ  is  concerned  a  few  general  facts  of 
significance  may  be  mentioned. 

It  was  formerly  thought  that  the  liver  varied  inversely 
as  the  size  of  the  animal  but  Magnan  (1)  and  others  have 
shown  that  the  matter  is  not  so  simple.  In  the  first  place 
if  there  be  an  actual  mathematical  formula  it  is  that  the 
liver  varies  in  size  inversely  as  the  surface  area  of  the 
body,  but  this  is  not  the  whole  story.  It  seems  that  the 
relation  of  size  of  the  organ  to  its  weight  is  not  constant 
and  that  it  is  better  to  judge  of  the  organic  capacity  by 
the  latter.  In  herbivorous  animals,  both  birds  and  mam- 
mals, the  liver  is  lightest  per  kilo  of  body  weight ;  next  in 
weight  are  in  order,  fisheaters,  meateaters,  insectivora, 
seedeaters,  fruiteaters  and  omnivora.  There  is  besides 
this  a  roughly  inverse  ratio  between  the  size  of  the  liver 
and  the  length  of  the  intestine  and  in  the  class  Aves 
inversely  as  the  size  of  the  lungs  also.  From  the  im- 
mediately foregoing  statements  it  is  apparent  that  a 
bewildering  variation  occurs  and  that  only  rough 
measurements  of  the  relative  volume  of  the  liver  are  avail- 
able. An  attempt  was  made  in  the  Marsupialia,  which 
present  all  the  variations  given,  to  discover  if  any  peculiar 
pathology  corresponded  with  the  above  groups ;  as  it  was 
fruitless,  no  change  from  our  zoological  treatment  will 
be  made. 

Lobar  arrangement  varies  from  the  relatively  simple 
double  avian  type  to  the  manifold  lobulations  of  the  seal 
or  the  marsupial  but  I  can  find  no  literature  to  indicate 
that  lobes  or  lobulations  have  a  direct  effect  upon  func- 
tions.    There  must  be  a  difference  of  blood  supply  for  in 

(1)  C.  R.  Soc.  de  Biologie,  Paris,  T.  73-526.  Bull.  Mus.  Hist.  Nat., 
Paris  Ann.,  1911,  492  et  seq. 


224  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

certain  infectious  diseases  like  enterohepatitis  and  amoe- 
biasis,  the  cystic  and  extreme  right  lobes  are  more  affected 
than  the  left  parts  of  the  organ.  In  the  bird  this  is  not 
so  difficult  to  follow  since  the  three  divisions  of  the  portal 
vein,  while  they  combine  at  times  in  an  ampulla  within  the 
hilum  of  the  liver,  seem  directed  to  certain  lobes,  that 
from  the  left  portal  seeming  to  point  toward  the  right 
side.  The  avian  portal  system  differs  from  the  mammalian 
in  having  a  large  branch  from  the  renal  area,  the  so-called 
renal-portal  system,  pass  to  the  liver,  and  by  having  a 
free  anastomosis  between  the  portal  area  and  the  caudal 
vena  cava  whereby  blood  from  the  pelvic  district  may 
pass  into  the  general  circulation  mthout  going  through 
the  liver.  There  is  no  unanimity  of  opinion  as  to  the  func- 
tion or  importance  of  this  connection  (2)  and  from 
the  data  collected  here  there  is  no  pecuhar  reno- 
hepatic  pathology. 

The  gall-bladder  is  not  a  constant  organ  in  either  mam- 
mals or  birds  and  indeed  it  may  be  absent  or  present  in 
very  closely  related  species  (Two-toed  Sloth  present, 
Three-toed  Sloth  absent).  When  present  in  mammals  it 
is  usually  a  dependent  bag  while  in  birds  it  commonly  lies 
upon  the  cystohepatic  duct,  between  the  liver  and  the 
last  curve  of  the  duodenum,  in  some  varieties  filling  from 
the  bottom,  the  inlet  being  guarded  by  a  valve.  This 
cystic  duct  in  nearly  all  birds,  comes  exclusively  from  the 
right  lobe  while  the  hepatic  duct,  with  which  the  cystic 
has  no  connection,  is  formed  by  combination  within  the 
liver  of  radicles  from  both  sides.  It  passes  to  the  duo- 
denum well  in  advance  of  the  cystic  duct,  in  some  birds, 
e.g.,  the  Struthiones,  very  near  the  pylorus,  that  is  on 
the  descending  limb  of  the  duodenal  loop.  By  this  means 
obstruction  to  the  biliary  stream  is  rendered  difficult.  The 
common  duct  combines  with  one  of  the  pancreatic  outlets 
in  most  mammals  but  the  abdominal  salivary  gland  in 

(2)  See  Woodland,  Proc.  London  Zool.  8oc.,  1906,  and  MacLeod, 
Chemical  and  Physiological  Medicine,  Chicago,  1923. 


THE  LIVER  225 

lower  animals  has  more  often  patent  separate  ducts  or 
multiple  ducts  than  it  does  in  man.  Birds  have  one  to 
four  pancreatic  ducts  separate  from  the  biliary  openings. 

The  gall-bladder  is  missing  in  most  varieties  of  the 
following  groups:  pigeons,  parrots,  wrens,  ostriches, 
rheas,  cuckoos,  toucans  among  the  birds ;  most  odd-toed 
ungTilates,  hyraces,  Indian  elephants,  all  deer,  peccaries, 
three-toed  sloth,  and  many  rodents.  The  varieties  lack- 
ing this  reservoir  are  herbivorous  in  the  main,  true 
carnivores  seeming  always  to  be  possessed  of  such  a 
structure.  Among  the  important  herbivorous  ungulates, 
Bovidae,  Tragulidas,  Camelidse  and  Suidae  have  this  bile 
reservoir  almost  without  exception.  Because  of  the 
interest  now  being  shown  in  the  pathology  of  the  gall- 
bladder and  its  passages  and  of  the  pancreas,  it  was  hoped 
that  evidence  of  definite  practical  value  for  human  pa- 
thology would  be  at  hand  in  our  study  if  we  divided  the 
animals  into  groups  Avith  and  without  a  bile  reservoir. 
The  result  is  not  unequivocal  but  worthy  of  note;  it  is 
discussed  on  pages  238  and  255. 

Microscopically  the  well  known  lobular  arrangement 
of  the  liver  is  rather  faithfully  carried  out  among  the 
mammals  albeit  the  most  systematic  and  complete  archi- 
tecture is  to  be  found  in  the  pig  while  the  marsupial  seems 
the  most  disorderly,  thus  resembling  the  avian  organ.  In 
the  latter  class  all  the  parts  are  indistinct,  the  cells  having 
an  unclear  outline,  the  tubules  being  intricately  wound 
and  the  interlobular  connective  tissue  being  scanty  and 
not  anastomosing  in  a  definite  framework.  The  intra- 
lobular reticulum  is  especially  difficult  to  detect.  Groups 
of  cells  are  often  found  at  portal  spaces ;  these  are  large 
and  small  mononuclears  and  granular  cells,  probably  of 
the  hematopoietic  system.  It  is  possible  that  blood 
formation  is  performed  in  the  liver  and  spleen  in  some 
adult  birds  but  such  a  function  is  denied  for  the 
mammal  except  under  very  unusual  conditions  of  bone 
marrow  atrophy.  \ 


226   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Glycogenic  and  fatty  conservation  is  a  function  pos- 
sessed by  both  zoological  classes  as  are  the  detoxicating 
and  bile-producing  powers.  However  it  is  highly  prob- 
able that  urea  and  creatin  in  metabolism  is  not  cared  for 
by  the  avian  liver  as  it  is  by  the  maimnalian,  judging  by 
the  researches  of  Paton  and  of  Richet. 

Fat  Deposits. 

The  care  of  fat  by  the  liver  is  very  well  shown  by 
examining  the  incidence  of  fatty  metamorphoses  through 
the  various  orders.  In  the  first  place  Mammalia  show  a 
slightly  higher  percentage  of  fatty  change  than  do  Aves 
and  should  show  a  greater  difference  were  it  not  for  the 
large  number  of  cases  in  two  orders  of  the  latter.  Among 
mammals,  lemurs,  rodents  and  marsupials  store  fat  in  the 
liver  more  than  other  orders  but  in,  the  second  and  third, 
it  is  chiefly  the  carnivorous  varieties  that  have  this 
property.  Just  why  the  slothful  herbivorous  lemurs 
should  be  first  on  the  list  is  not  evident  especially  since 
the  grain-eating  UngTilata  are  least  apt  to  present  fatty 
livers.  With  tliis  exception,  mammals  with  plentifully 
available  hydrocarbons  in  their  diet  are  most  apt  to  show 
its  deposit  in  the  organ  under  discussion.  Among  the 
birds  the  gallinaceous  varieties  stand  far  ahead  of  aU 
others,  the  passerines  following  next.  Galli  show  the  con- 
dition in  association  with  acute  infections,  chronic  dis- 
eases and  in  health.  Unless  there  be  distinct  reason  for 
it  at  autopsy,  it  may  almost  always  be  said  to  be  normal^ 
Passeres,  especially  the  smaller  forms,  frequently  come 
to  autopsy  with  such  excessively  large  livers,  and 
indeed  with  a  very  large  pad  of  abdominal  fat,  and 
nothing  else,  that  one  is  compelled  to  look  upon  this  over- 
burdened organ  as  incapacitated  by  the  deposit.  These 
two  orders  increase  the  percentage  value  for  the  birds. 
Striges,  Anseres,  and  Accipitres  also  show  a  good  number 
of  cases  but  there  is  among  the  Aves  no  such  clear  rela- 


THE  LIVER  227 

tionsliip  between  food  fat  and  fat  infiltration  as  may  be 
found  in  the  Mammalia. 

Amyloid  Deposits. 

Amyloid  deposit  is  reported  with  reasonable  frequency 
in  domesticated  animals,  causing  in  them  a  fairly  definite 
entity,  being  as  usual  related  to  the  effects  of  long  con- 
tinued or  repeated  infectious  disease.  Wild  animals  suf- 
fer from  this  condition  but  rarely  and  therefore  to  our 
few  cases  will  be  given  a  short  discussion  separately.  An 
Indian  Paradoxure  {Paradoxurus  niger)  had  patches  of 
amyloid  irregularly  distributed  through  the  organ.  The 
animal  had  a  carcinoma  of  the  head  of  the  pancreas,  an 
obstructive  biliary  cirrhosis  in  a  state  of  atrophy  and 
a  chronic  nephritis  with  arteriosclerosis.  There  was 
nothing  peculiar  about  the  distribution  of  the  deposit  as 
there  was  in  the  next  case,  a  Badger  (Meles  meles)  where 
amyloid  was  found  around  the  interlobular  vessels  and 
extending  in  the  lobules  along  their  canaliculi.  This 
latter  case  seemed  wdthout  cause  and  we  have  considered 
it  a  primary  amyloidosis,  the  spleen,  heart  muscle,  kid- 
neys, intestines  and  other  structures  being  affected.  ( See 
Fig.  7.)  A  third  mammalian  case  concerned  a  Dasyure 
(Dasyurus  viverrinus)  which  showed  distinct  intralobu- 
lar collections.  Its  cause  was  a  chronic  suppurative 
process  in  the  jaw  bone. 

.  Avian  livers  are  somewhat  more  prone  to  show  amy- 
loid deposits,  eight  cases  being  on  record.  Four  occurred 
in  the  Passeres,  one  each  in  ColumbEe  and  Impennes  and 
two  in  Anseres.  Three  were  associated  with  chronic 
infectious  disease  and  two  with  well  established  nematode 
parasitism.  The  remaining  three,  classed  as  primary, 
were  not  related  to  any  other  lesions,  in  two  the  amyloid 
liver  being  the  only  finding. 

The  next  abnormal  deposition  related  to  the  physiology 
of  the  organ  is  blood  pigmentation.  Normally  hemic  pig- 
ment is  dispensed  with  veiy  rapidly  but  under  unnatural 


228  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

conditions  it  accumulates.  In  only  one  order  is  there  any- 
noteworthy  percentage  of  hemosiderosis,  the  carnivores, 
the  remainder  showing  a  very  trifling  incidence. 

Degenerations. 

Going  further  into  the  physicochemical  alterations  of 
the  liver  brings  us  to  consideration  of  those  changes 
known  as  degenerations — parenchymatous,  fatty,  hy- 
dropic, hyaline,  all  of  which  we  shall  group  under  one 
heading.  They  occur  in  a  great  variety  of  conditions  and 
do  not  appear  to  be  specific,  nor  as  the  records  are 
analyzed  do  they  appear  to  occur  preeminently  in  any  one 
disease  of  the  lower  animals.  The  percentages  are  how- 
ever higher  for  orders  and  families  whose  diet  contains 
relatively  more  protein,  carnivores,  the  higher  mar- 
supials, accipitrine,  and  wading  birds. 

Acute  Atrophy. 

,  A  very  important  degenerative  disease  of  the  liver  is 
aciite  yellow  atrophy  or,  better  expressed,  acute  degenera- 
tive atrophy  for  it  is  a  total  destruction  of  the  w^hole  or 
large  parts  of  the  parenchyma.  It  is  apparently  toxic  in 
origin  being  related  to  the  toxemias  of  pregnancy,  to 
certain  organic  and  inorganic  soluble  poisons ;  some  cases 
arise  without  discoverable  cause.  We  have  seen  no  cases 
in  the  mammal  but  two  in  birds.  Both  were  females,  one  in 
active  ovulation,  while  the  other  had  no  related  pathology 
and  the  condition  of  the  ovaries  could  not  be  deter- 
mined since  they  had  been  destroyed  after  death  by  rats. 
The  macroscopic  and  minute  anatomy  offers  nothing  new. 
Jaundice  was  present  but  not  intense.  , 

Hepatitis. 

True  inflammatory  lesions  are  to  be  defined  as  some 
form  of  parenchymatous  change  to  which  are  added  con- 
gestion, infiltration  of  round  or  polynuclear  cells,  stagna- 
tion in  the  bile  ducts  or  perhaps  actual  degeneration  of 


THE  LIVER  229 

their  lining  cells.  It  seems  necessary  to  stipulate  these 
things  because  in  the  chronic  forms,  usually  called  cir- 
rhosis, it  is  necessary  to  have  all  of  them,  plus  efforts  at 
regeneration,  in  order  to  determine  it  as  a  chronic  pro- 
gressive process.  Acute  hepatitis  is  a  rare  condition  in 
mammals  except  when  it  is  combined  with  septicemia  or 
severe  enteritis.  In  birds  on  the  other  hand  the  liver  is, 
aside  from  the  intestinal  wall,  perhaps  the  most  frequent 
seat  of  pathology  in  the  abdomen.  This  is  because  of  its 
almost  constant  involvement  in  infective  enteritis,  and  in 
such  conditions  as  fowl  cholera,  fowl  typhoid,  coccidiosis 
and  cecal  amcebiasis,  all  of  which  we  have  sporadically. 
When  one  searches  for  special  distribution  among  the 
orders,  only  one  of  them  stands  out  as  having  a  high  per- 
centage, the  Galli,  an  order  which  seems  to  have  a  very 
vulnerable  liver. 

The  macroscopic  anatomy  of  hepatitis  in  birds  is 
peculiar  in  showing  a  definite  swelling  with  spots  of  gray 
or  yellow  color,  sometimes  coalescing  to  form  irregailar 
areas.  These  are  much  more  definite  than  in  the  mam- 
malian organ  where  swelling  and  hemorrhage  are  the 
commoner  findings.  These  pale  spots  are  of  two  origins. 
They  may  be  focal  necroses  of  the  hepatic  cells,  with  or 
without  circumferential  congestion  or  hemorrhage  to 
make  them  stand  out.  In  amoebic,  coccidial  and  typhoid 
livers  such  is  the  type  of  change.  In  septicemia  and 
cholera,  the  mottlings  are  made  up  of  increased  inter- 
stitial mononuclear  areas,  mth  blood  cells  and  shadow 
cells  numerously  present.  I  have  seen  what  was  in  all 
probability  a  stage  of  repair  after  both  these  kinds  of 
change.  In  the  former,  regeneration  seemed  to  take  place 
from  adjoining  liver  cells,  there  being  in  the  section  no 
evidence  of  increased  bile  ducts  to  make  new  hepatic  cells. 
It  seemed  also  that  phagocytes  were  derived  from  blood 
cells  and  not  from  Kupffer's  cells.  In  the  infiltrative 
lesion  disappearance  of  the  liver  cells  from  the  groups 
leaving  compressed  and  deeply  granular  remnants  was  all 


230  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

that  could  be  determined.     Regeneration  seemed  to  be 
progressing  in  the  manner  just  outlined,  i 

Neckoses. 

,^  The  degenerative  and  infiltrative  areas  of  acute  hepa- 
titis are  simulated  by  focal  necroses  in  livers  not  the  seat 
of  a  general  hepatitis  from  which  they  can  be  differen- 
tiated only  by  the  microscope.  These  small  areas  of 
local  tissue  death  are  quite  common  in  all  pathological 
processes  but  are  most  common  in  the  liver,  possibly 
because  of  its  exposure  to  toxins  from  the  intestine.  Their 
exact  origin  is  not  determined,  various  explanations 
being  given.  The  somewhat  distinct  distribution  in  mam- 
mals versus  that  in  birds  may  help  in  the  final  decision. 
In  the  former,  focal  necroses  are  more  often  encountered 
midway  in  the  anatomic  lobule  and  around  the  central 
vein  whereas  a  perivascular  location  seems  the  usual 
position  in  the  bird. 

Massive  necroses  of  the  liver  may  be  of  considerable 
importance  in  veterinary  medicine.  They  take  their  origin 
in  several  different  ways.  The  commonest  in  our  records 
are  those  due  to  cecal  coccidiosis  and  amoebiasis  (quail 
disease  and  blackhead)  while  from  the  primary  seat  of 
these  two  infectious  diseases,  the  cecum,  may  originate 
the  virus  of  non-specific  hepatic  necroses.  We  have 
observed  several  birds,  passerine,  psittacine  and  galli- 
naceous, which  at  autopsy  showed  a  distention  of  the 
cloaca,  ceca,  and  lower  small  intestine  with  urates  and 
slime  but  no  mural  inflammation  and  a  large  area  of 
necrosis  in  the  liver.  This  suggests  perhaps  a  **  white 
diarrhoea  "  but  it  did  not  occur  in  epizootics  and  other 
morbid  anatomy  of  this  specific  disease  was  absent.  These 
frequent  instances  of  association  between  the  colonic  area 
and  the  liver  seem  to  suggest  the  transfer  of  necrotizing 
organisms,  just  as  amcebae  travel,  and  to  indicate  measures 
to  clean  out  the  tract  when  birds  become  ''  plastered." 
Massive  necroses  also  arise  from  mould  disease,  and  from 


THE  LIVER  231 

infection  with  the  necrosis  bacillus,  emanating  from 
nearby  infectious  foci,  or  via  the  normal  passageways 
from  the  intestine.  Massive  areas  of  degeneration  may 
form  by  the  coalescing  of  numerous  foci,  in  any  septi- 
cemic disease. 

Abscess. 

In  man,  amoebae,  flukes,  cestodes  and  biliary  tract  infec- 
tion are  the  commonest  causes  of  purulent  collections 
within  the  liver.  In  the  lower  mammals  parasites  play 
practically  a  solitary  role  at  least  as  the  major  influence 
in  localizing  the  collection,  bacteria  from  the  intestine 
doing  the  rest.  We  have  one  case  of  massive  abscess  in 
a  porcupine  suffering  with  septic  pneumonia,  the  sup- 
puration in  the  liver  being  due  to  the  colon  bacillus,  the 
general  septicemia  probably  being  from  distemper. 
Monkeys  have  shown  more  abscesses  than  any  other 
order,  three  being  observed.  One  was  due  to  infestation 
with  trichocephalus  which  had  apparently  penetrated 
from  the  colonic  wall  into  the  liver  through  adhesions 
formed  between  these  two  structures.  Another  seems 
certainly  amoebic  but  these  protozoa  could  not  be  found, 
while  the  third  followed  an  ulcerative  enterocolitis  of 
unknown  cause.  Two  cats  were  seen  with  parasitic 
abscesses ;  one  harbored  Distoma  or  ClonorcJiis  sinensis, 
the  other  a  nematode  of  ascaris  type. 

The  topographic  distribution  of  these  six  hepatic 
abscesses  was  interesting.  The  position  of  the  abscess  is 
not  mentioned  in  one  case  but  of  the  remaining  five  three 
were  entirely  in  the  right  lobe,  one  had  the  major  lesion  on 
the  right  side  and  smaller  separate  abscesses  spread  over 
the  organ,  and  one  with  about  equal  distribution  in  all 
lobes.    All  three  confined  to  the  right  side  were  solitary. 

Abscesses  of  considerable  size  are  not  met  with  in  the 
bird  as  in  the  mammal  perhaps  because  the  former  does 
not  form  real  pus,  necroses  developing  instead. 


232   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Congestion  of  the  liver  is  a  matter  of  small  importance 
from  the  standpoint  of  pathology  unless  it  be  of  sufficient 
duration  to  cause  cyanotic  atrophy  and  induration.  How- 
ever the  facts  that  congestion  of  this  organ  occurs  three 
times  as  often  in  the  mammal  as  in  the  bird  and  that 
vascular  cirrhosis  has  not  been  seen  in  the  latter  class, 
are  interesting  and  noteworthy.  [In  addition  ascites  of 
hepatic  origin  has  not  been  seen  in  the  birds.  The  expla- 
nation for  this  lies  in  the  rich  anastomosis  between  the 
intestinal  area  and  the  caudal  vena  cava  so  that  the  blood 
does  not  have  to  pass  through  the  liver  to  reach  the  heart. 
This  arrangement  would  reduce  the  back  pressure  in 
passive  congestion  and  relieve  the  liver  in  the  congestion 
due  to  toxic  or  inflammatory  distention  of  small  vessels.  \, 

ClERHOSIS. 

The  chronic  inflammations  or  so-called  cirrhoses  of  the 
liver  have  been  subjected  to  a  great  deal  of  study  and 
many  theories  have  been  expounded  as  to  their  cause  and 
classification.  Here  is  not  the  place  to  discuss  the  aca- 
demic question  of  nomenclature  but  rather  to  adopt  an 
acceptable  worldng  classification  and  to  analyze  our 
material  thereon.  A  cirrhosis  is  a  chronic  inflammation 
of  the  liver  indicated  by  increased  connective  tissue  with 
evidences  of  degeneration  and  attempts  at  regeneration 
on  the  part  of  the  hepatic  cells.  Certain  cases  of  increased 
connective  framework  fail  to  show  the  last  two  features 
and,  since  they  must  be  grouped  near  the  cirrhoses  because 
of  the  prominence  of  connective  tissue,  they  are  called 
fibroses,  perilobular  in  type.  Among  the  instances  carry- 
ing out  the  full  stipulations  are  livers  with  evidence 
of  a  perivascular  fibrosis  and  obstruction,  to  which  are 
added  degeneration  and  regeneration  of  the  lobular 
margins;  such  are  portal,  cierhoses  in  human  medicine 
associated  with  passive  congestion  in  the  intestinal  area, 
and  ascites.  In  a  second  variety,  fibrosis  seems  to  succeed 
upon  obstruction  to  the  biliary  lumina  or  upon  peribiliary 


THE  LIVER  233 

inflammation,  biliary  cierhoses.  The  effects  of  this  are  to 
dam  back  bile  with  the  production  of  varying  degrees  of 
jaundice  and  for  the  inflammation  to  spread  into  the 
lobules,  thus  distorting  their  internal  architecture;  this 
form  is  therefore  unlike  portal  cirrhosis  which  alters  the 
size  and  shape  of  lobules  as  a  whole.  Fatty  change  is 
very  prominent  in  certain  cases  and  it  has  been  a  custom, 
perhaps  without  warrant,  to  put  such  livers  into  a 
separate  group.  It  may  be  that  they  represent  a  different 
chemical  process.  /When  there  exists  for  a  long  time  a 
venous  stasis  in  the  liver,  necrosis  is  apt  to  occur  in  the 
cells  subjected  to  pressure  and  the  absence  of  fresh  blood. 
This  gives  rise  to  a  '*  nutmeg  "  liver  upon  which  may 
succeed  a  definite  perivenous  fibrosis. 

This  then  is  a  working  classification  of  the  hepatic 
cirrhoses.  Perhaps  many  slightly  differing  varieties 
might  be  constructed  but  this  grouping  will  permit  com- 
parison and  contrast  with  human  cases,  and  with  instances 
in  the  various  orders.  Because  of  the  relatively  small 
total,  thirty-two,  it  is  perhaps  unwise  to  attempt  any  con- 
clusions as  to  distribution  but  it  is  certainly  noteworthy 
that  twenty-six  occurred  in  mammals.  This  means  1.6 
per  cent,  in  mammalian  autopsies  against  .2  per  cent,  in 
avian.  Among  the  former  class  the  carnivores  stand  at 
the  head  of  the  list,  followed  in  order  by  the  marsupials, 
ungulates,  primates,  and  rodents. 

Carnivora  have  sho^vn  a  few  typical  portal  cirrhoses 
from  a  pathological  standpoint  but  only  one,  in  a  badger 
{Taxidea  taxus),  was  combined  with  the  classical  picture 
of  intestinal  hyperemia  and  ascites.  Two  of  the  cases  were 
combined  with  chronic  enteritis  which  may,  of  course, 
have  been  secondary  but  there  was  also  a  hyperplasia  of 
the  spleen  which  bespoke  some  grade  of  infection.  None 
of  the  four  showed  involvement  of  the  biliary  tract.  One 
animal,  a  skunk  {Mephitis  mesomelas),  was  jaundiced; 
it  had  anemia,  nephritis  and  enlarged  spleen  but  no 
intestinal  inflammation;  perhaps  the  associated  anemia 
16 


234  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

may  have  been  responsible  for  the  pigmentation.  Biliary- 
cirrhosis  occurred  in  two  Carnivora,  in  both  associated 
with  enlarged  spleen  and  nephritis.  One  showed  jaundice 
and  the  other,  with  a  huge  liver  from  congestion  and 
interstitial  infiltration,  had  a  small  ascites.  Fatty  cir- 
rhosis was  diagnosed  in  a  raccoon  but  this  is  viewed  with 
some  reservation  because  this  animal  easily  stores  fat 
and  in  this  case  it  may  not  have  been  a  part  of  the  process. 
In  none  of  the  foregoing  cases  did  parasitism  enter  into 
the  causation  of  the  change  and  I  shall  always  specify 
when  such  a  factor  was  probable.  The  only  vascular 
cirrhosis  in  our  records  occurred  in  a  Gray  Wolf  incident 
to  a  longstanding  myocarditis  (Gray  Wolf,  Cams  lupus 
mexicanus,  Myocarditis,  Adenomatoid  goitre,  Chronic 
gastroenteritis.  Vascular  cirrhosis  of  liver,  Subacute  dif- 
fuse nephritis.  Edema  of  lungs,  pericardium,  and  peri- 
toneum). Two  examples  of  perilobular  fibrosis  appeared 
in  this  order,  a  raccoon  {Procyon  lotor)  and  a  paradoxure 
{Trichosurus  vulpecular  vulpecular).  The  only  note- 
worthy feature  was,  in  the  former,  a  very  marked  biliary 
stasis  on  the  lobular  margins  and  in  the  connective  tissue ; 
this  animal  was  not  jaundiced. 

Ungulata  are  normally  well  supplied  with  definite 
interlobular  strands  which,  in  a  few  varieties,  completely 
encircle  the  lobule  but  always  show  as  clear  fibrous  septa 
going  out  from  the  portal  areas.  This  richness  of  con- 
nective tissue  renders  more  difficult  a  decision  of  increase 
so  that  unequivocal  degenerations  and  regenerations 
with  inflammatory  changes  have  been  demanded  as  cri- 
teria for  cirrhosis.  It  has  been  recognized  that  cattle  get 
a  definite  increase  in  their  interstitial  tissue  without 
serious  reaction  in  the  parenchyma.  With  the  knowledge 
of  these  facts  in  mind  it  has  been  possible  to  detect  two 
distinct  portal  cirrhoses,  two  biliary  cirrhoses  and  three 
perilobular  fibroses.  It  is  however  evident  by  examining 
the  rest  of  the  autopsy  notes  that  the  chronic  inflam- 
mations have  had  with  one  exception,  little  influence  on 


THE  LIVER  235 

the  animal's  life  and  death  and  the  associated  pathology- 
is  not  instructive  in  etiology.  One  old  deer  with  the 
definite  portal  type  had  ascites  and  intestinal  hyperemia 
which  hastened  his  end. 

The  tjrpe  of  cirrhosis  in  the  marsupial  is  progressively 
inflammatory  and  of  the  biliary  variety.  In  two  of  the 
three  cases  there  was  active  infection  somewhere  in  the 
body,  one  a  long  continued  streptothricosis,  the  other  and 
more  important  a  choledochitis  with  involvement  of  the 
pancreatic  head.  The  third  case  showed  a  nephritis  and 
a  pericholedochitis  and  pericholecystitis.  In  all  three 
there  was  definite  evidence  of  biliary  obstruction  within 
the  liver  and  in  the  occurrence  of  general  jaundice. 

Monkeys  have  presented  one  portal,  two  biliary,  and 
one  perilobular  cirrhoses.  The  London  Garden  reports  a 
cirrhosis  mth  gall  stones  in  a  Chimpanzee.  The  case  of 
the  Barbary  Ape  is  so  good  that  it  is  quoted  in  brief. 

Barbary  Ape  6  {Macacus  innuus).  Found  dead.  Never  known  to 
be  sick.  On  exhibition  nine  years.  Acute  dilatation  of  stomach.  Acute 
gastritis.  Portal  cirrhosis  of  liver.  Acute  parenchymatous  nephritis. 
Chronic  passive  congestion  of  lungs.  Chronic  splenitis  and  perisplenitis. 
Ascites.  Mild  passive  congestion  of  abdominal  circulation.  On  open- 
ing the  abdomen  a  dilated  stomach  occupies  most  of  the  anterior  part, 
displacing  the  intestines  downward  and  backward.  The  upper  lobes 
of  both  lungs  are  uniformly  deep  red,  soft,  collapsed,  subcrepitant.  Sub- 
clavian vessels — veins  distended  with  red  clot,  arteries  with  small  amount 
of  chicken  fat  clot.  The  heart  is  dilated  on  the  right  side,  filled  with 
currant  jelly  clot.  The  liver  is  small,  surface  hobnailed,  edges  rough, 
consistency  tough,  color  brown.  Section  surface  glistening,  moist, 
granular  and  opaque,  mottled  by  iiTegular  brown  areas  separated  by 
paler  brown  intercommunicating  bands.  Gall-bladder  is  small,  contains 
viscid  yellow  bile  and  duct  is  patulous.  Areolar  tissue  about  the  bile 
ducts  is  thick  and  opaque,  the  duct  wall  itself  is  thick  and  yellow.  Gall- 
bladder tightly  attached  to  capsule  of  liver.  Spleen  is  slightly  enlarged, 
soft  and  tough.  Capsule  is  smooth,  opaque  and  thickened  on  gastric 
surface.  The  trabeculee  are  prominent,  pulp  mottled  gray-red,  few 
recent  hemorrhages.  Capsule  of  the  kidneys  is  smooth,  strips  easily 
leaving  a  smooth  bro-\vn  surface  with  dilated  vessels.  Organ  is  soft. 
Section  surface  is  glistening,  striae  wide  and  indistinct,  glomeruli  faintly 
visible.  Microscopic  section  of  liver  shows  high  grade  of  fibrosis 
almost  entirely  confined  to  portal  areas  with  a  marked  increase  in  bile 
ducts  although  no  place  is  found  where  these  bile  ducts  are  running 


236  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

into  lobules  suggesting  attempt  at  regeneration.  Liver  cells  show  high- 
grade  of  fatty  degeneration  in  some  places,  whole  lobules  being  necrotic. 
There  is  no  pigmentation  and  connective  tissue  is  fairly  rich  in  cells. 
Fibrosis  quite  well  advanced.  Cells  about  equally  fibroblasts,  round 
cells  and  polynuclears.  Bile  ducts  very  Avell  preserved  and  cellular 
infiltrate  rather  less  directly  around  them  than  at  other  parts  of  con- 
nective tissue.  The  section  of  kidney  shows  moderate  congestion,  granu- 
lar and  vacuolar  degeneration  of  epithelium  generally  distributed  except 
in  proximal  tubules  where  there  is  swelling  and  desquamation.  Detritus 
present  in  tubules  and  capsular  spaces.     Tufts  swollen. 

The  biliary  forms  of  Primates  were  associated  in  one 
case  with  an  undetermined  parasite  in  the  bile  channels,  in 
the  other  with  tuberculosis  and  chronic  enteritis.  In  all 
the  cases  the  relative  inconspicuousness  of  bile  in  ducts 
or  in  cells  is  worthy  of  mention.  The  perilobular  fibrosis 
in  a  small  cebus  was  trifling  in  extent  but  was  associated 
with  considerable  round  cell  infiltration  in  isolated  areas ; 
there  was  also  nephritis,  splenitis,  and  enteritis. 

The  only  representative  of  the  rodents  is  a  capybara 
{Hydrochoerus  hydrochoRrus),  their  largest  variety.  This 
case  was  originally  described  as  a  typical  Laennec  or 
Pictou  cirrhosis  but  I  now  class  it  as  a  portal  form.  The 
distinct  insular  arrangement  of  the  lobules,  the  failure  of 
involvement  of  the  bile  channels  and  the  ascites  are 
reasons  for  the  present  decision.  The  animal  suffered 
also  from  tuberculosis  (not  in  liver)  and  myocarditis. 

The  Indian  Elephant,  ''Bolivar"  {Elephas  indicus), 
an  old  specimen,  is  the  only  member  of  his  order  to  show 
cirrhosis.  It  may  be  considered  as  a  senile  process 
in  part  but  the  extreme  distortion  and  compression  of 
the  lobules  press  the  conclusion  that  it  was  a  pro- 
gressive inflammation. 

Aves  fail  to  show  lesions  which  could  be  called  portal 
cirrhosis,  five  of  their  six  cases  being  biliary  and  one  fatty 
with  sigiis  of  continued  infection.  The  macroscopic 
anatomy  of  the  avian  liver  with  chronic  fibrosing  hepatitis 
is  fairly  uniform  and  suggestive.  In  the  first  place  it  is 
grossly  nodular,  lumpy,  not  finely  granular  or  ' '  hob- 


Fig.    18. — PORTAL  CIRRHOSIS  OF  LIVER  IN  ATROPHIC  STAGE 
APE   (MACACUS  INNUUSh      THE  DILATATION  OF  THE  STOMACH   AL: 
IN   PHOTOGRAPH. 


THE  LIVER  237 

nailed."  The  sensation  to  the  finger  is  resilient  rather 
than  tough.  The  color  is  variable  but  green  and  dull 
purple  are  the  usual  shades.  On  section  no  peculiarities 
present  themselves  unless  it  be  that  one  can  find  pale  spots 
on  a  dark  background,  which  may  correspond  to  the  mam- 
malian connective  tissue  strands.  Microscopically  the 
increase  of  cellular  groups  at  portal  spaces  and  the  exten- 
sive growth  of  connective  tissue  between  the  liver  columns 
are  the  noteworthy  features.  There  is  nothing  in  mam- 
malian cirrhoses  to  compare  with  the  intralobular  growth 
of  fibres  in  birds.  There  is  of  course  no  regularity  so 
that  the  degree  of  replacement  or  necrosis  of  parenchyma 
is  hard  to  estimate.  Bile  ducts  do  not  proliferate  but 
seem,  once  obstructed  and  surrounded,  to  succumb  to  the 
inflammation,  The  six  cases  in  birds  are :  Psittaci,  3,  Galli, 
Anseres,  Struthiones  each  one.  The  cases  in  the  last  two 
orders  were  associated  with  parasites,  to  which  bacteria 
or  toxin  may  have  been  added.  It  is  interesting  to  note 
that  the  two  frankly  progressive  obstructive  biliary  cases 
in  the  parrots  showed  general  jaundice. 

It  was  formerly  customary  in  many  quarters  to  speak 
of  atrophic  and  hypertrophic  cirrhosis.  Now  it  is  gen- 
erally thought  that  any  form  will  be  large  or  small  as 
growth  and  regeneration  on  the  one  hand,  or  contraction, 
atrophy  and  degeneration  on  the  other,  may  be  predomi- 
nant at  the  time  the  organ  is  seen.  It  is  perhaps  mislead- 
ing to  judge  by  our  notes  of  what  happens,  but  it  is 
curious  that  in  the  thirty-two  cases,  the  pathologist  could 
state  only  in  seventeen  instances  that  the  liver  was  larger 
or  smaller  than  normal.  This  means  therefore  that  the 
liver  of  cirrhosis  need  not  deviate  greatly  from  its 
customary  size.  Nine  of  the  seventeen  times  the  organ 
was  considered  smaller  than  normal,  eight  times  it  was 
greater.  These  variations  did  not  strictly  correspond  to 
type,  but  the  portal  form,  frequently  called  atrophic,  was 
more  often  small  than  was  the  biliary  form. 


238   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Gastrointestinal  disease  accompanied  cirrhosis  in 
fourteen  instances.  Nephritis  was  present  nineteen  times. 
The  spleen  was  enlarged  six  times,  in  all  of  which  definite 
evidence  of  infection  existed  in  the  body.  Choledochitis 
existed  four  times,  twice  with  biliary  cirrhosis,  twice  ^\dth 
perilobular  fibrosis ;  cholecystitis  existed  twice,  once  in  a 
monkey,  and  once  in  a  bird  Avith  parasites.  Pancreatitis 
was  seen  in  three  biliary  cirrhoses  and  once  in  a  peri- 
lobular fibrosis. 

The  relation  of  the  existence  of  cirrhosis  to  the 
presence  of  a  gall-bladder  is  interesting.  Among  the 
thirty-two  animals  twenty-one  have  gall-bladders,  eleven 
have  not.  The  exact  number  of  animals  in  our  whole  list 
with  and  without  this  structure,  unfortunately  cannot  be 
given  with  exactness.  As  nearly  as  I  can  figure  it  out,  six- 
teen per  cent,  of  our  animal  posts  have  been  on  varieties 
without  a  gall-bladder,  eighty-four  per  cent,  with  it.  This 
would  make  the  absence  of  this  reservoir  a  factor  favoring 
the  development  of  cirrhosis  since  one-third  of  the  cir- 
rhoses are  in  groups  devoid  of  this  bag,  yet  these  same 
groups  supplied  only  one-sixth  of  the  total  postmortems. 

Gall  Stones. 

Our  experience  wdth  concrements  in  the  biliary  system 
is  limited  to  six  cases  which  can  be  detailed  in  brief. 

American  Beaver  9  {Castor  canadensis)  showed  a  soft  purplish 
liver  with  groups  of  tortuous  yellow  lines;  these  prove  to  be  groups 
of  hepaticola  with  fatty  degeneration  around  them,  but  successful 
regeneration  is  going  on ;  bile  ducts  are  not  seriously  involved  over  any 
great  part  of  the  organ ;  the  bladder  is  distended  greatly  with  thin,  yel- 
low-green fluid ;  duct  is  not  patulous ;  common  duct  narrowed  at  middle 
and  above  this  constriction  lies  a  small  concrement;  bladder  contains 
two  large  and  several  small  pale  yellow-green  friable  stones;  mucosa 
injected  and  covered  with  mucopus;  the  pancreas  is  not  affected. 

American  Beaver  6  {Castor  canadensis)  shows  a  slight  bile  obstruc- 
tion and  pigmentation  through  the  liver  but  no  pus  or  cirrhosis ;  bladder 
is  collapsed  containing  only  a  little  limpid  brown  fluid ;  wall  is  slightly 
roughened  but  not  opaque ;  there  is  a  blue-black  stone  1.5  x  1  cm.  free 
in  the  cavity;  duct  patulous;  pancreas  and  intestine  not  affected. 

Brant  Goose  6  {Branta  bernicla  glaucogastra)  liver  shows  slight 
fatty  change;  bladder  much  distended,  contains  twenty-six  small,  quite 


THE  LIVER  239 

hard,  greenish  stones;  one  is  impacted  in  the  cystic  duct  which  is 
not  patulous. 

Pigtailed  Macaque  6  (Macacus  nemestrinus)  shows  a  normal  liver; 
bladder  contains  a  small  black  concrement,  very  hard,  no  cystitis. 

Polar  Bear  ?  {Ursus  maritimus)  showed  a  chronic  cholecystitis  and 
cholangitis,  the  stone  (?)  in  this  case  consisting  of  a  solitary,  black, 
friable  mass,  six  mm.  in  diameter. 

Mongoose  Lemur  6  (Lemitr  mow^ro^)  showed  a  normal  liver;  bladder 
of  about  normal  size  but  the  duct  can  be  forced  only  by  considerable 
pressure ;  there  is  a  small  stone  and  a  granule  in  the  tortuous  cystic  duct ; 
no  cholecystitis. 

The  specimens  that  are  preserved  show  these  to  be 
chiefly  inspissated  bile,  those  from  the  first  beaver  and 
the  goose  being  the  only  ones  to  rise  to  the  dignity  of  gall 
stones ;  it  would  seem  that  there  was  plenty  of  opportunity 
for  calculi  to  form  in  the  bladder  of  this  beaver.  In 
no  case  is  there  a  cholangitis  or  cirrhosis  dependent 
upon  cholelithiasis. 

While  stones  have  been  showTi  as  infrequent  there  is 
a  condition  of  the  bile  which  may  be  quite  important.  In 
Passeres,  Accipitres,  and  Striges  one  frequently  sees  a 
very  dense  inspissation  of  the  bile  both  in  the  cystic  area 
and  in  the  lesser  independent  bile  duct.  This  need  not 
be,  indeed  usually  is  not,  associated  with  hepatitis  or 
cholecystitis.  There  is  no  one  thing  more  common  than 
another  in  relation  with  it  but  the  diagnoses  most  often 
made  are  enteritis,  distention  of  the  proventricle  and 
gizzard,  and  constipation. 

Inflammation  of  the  Biliaky  System. 

>^  The  biliary  tract  from  its  origin  in  fine  intrahepatic 
radicles  to  the  bladder  and  to  the  end  of  the  common  or 
intestinal  ducts  is  the  seat  of  many  inflammations  both 
acute  and  chronic,  but  since  they  are  supposed  to  lead  to 
damage  to  the  liver  and  pancreas  and  to  the  production  of 
gall  stones,  it  is  well  to  consider  the  system  as  a  whole. 
As  a  matter  of  fact  separate  analyses  of  cholangitis, 
choledochitis  and  cholecystitis  do  not  reveal  different 
figures  for  each  or  for  different  orders.  The  vulnerability 


240   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

of  this  tract  is  found  to  be  directly  as  the  percentage  of 
cirrhosis,  to  wit,  the  carnivores  stand  first,  then  the  mar- 
supials, ungulates.  Primates  and  rodents  |among  the  birds 
the  order  is  Accipitres,  Anseres,  Struthiones,  Psittaci,  and 
Galli.)  It  is  difficult  in  most  instances  to  evaluate  the 
various  possible  etiological  factors,  but,  due  caution  being 
exercised,;gastrointestinal  inflammation  could  be  held  re- 
sponsible in  seventeen  of  the  total  of  fifty  cases.  In  twelve 
of  the  seventeen  this  process  was  wholly  or  largely  in 
the  duodenum.) (The  next  factor  was  general  infection,  at 
the  head  of  wliich  pneumonia  and  ' '  distemper  ' '  occupied 
about  equal  places.  In  marsupials,  the  streptococcal  and 
streptothrical  infections  to  which  these  animals  are  sus- 
ceptible, was  the  prime  factor.  This  group  almost  always 
has  definite  signs  of  stasis  both  in  the  liver  and,  as  indi- 
cated by  jaundice,  in  the  general  tissues.  Pancreatitis 
was  present  in  seven  of  the  fifty  cases  and  in  five  of  the 
seven,  enteritis  was  also  found.  Common  duct  stones 
were  not  observed.  I  shall  have  sometMng  to  say 
about  pericholangitis  and  pericystitis  under  the  head 
of  pancreatitis. 

TUMOKS. 

The  liver  presents  a  good  share  of  the  tumors  appear- 
ing in  solid  viscera  but,  with  the  exception  of  a  few  points, 
they  offer  little  of  interest.  In  the  first  place  three  angio- 
mata  have  been  seen  and  while  they  may  not  be  tumors  in 
the  accepted  sense  of  the  word,  may  be  considered  briefly. 
A  single  cavernous  angioma  was  seen  in  a  goose.  It 
occupied  a  large  part  of  the  right  lobe  but  did  not  seem 
to  affect  mechanically  the  fmiction  of  the  organ  since  con- 
ditions wholly  foreign  to  the  liver  were  the  cause  of  death^ 
A  leopard  presented  several  small  groups  of  telangiec- 
tatic angiomata  lying  mostly  at  portal  spaces,  a  few  also 
under  the  capsule.  The  liver  of  a  thrush  was  likewise 
scatteringly  beset  with  small  angiomata.     The  original 


THE  LIVER  241 

notes  and  recent  examination  do  not  reveal  parasites  or 
perivascular  sarcomatous  change. 

:  Simple  adenomata  were  observed  in  a  woodchuck 
{Arctomys  monax).  This  diagnosis  is  made  with  the 
appreciation  that  nodular  regeneration  of  the  liver  after 
damage  and  in  cirrhosis  sometimes  suggests  tumor,  but 
with  adenomata  an  increase  of  supporting  framework 
may  occur.  The  liver  of  this  animal  presented  numerous 
.3  to  10.  cm.  irregularly  spherical,  encapsulated,  firm  or 
slightly  resilient,  bro^\^l  masses  which  under  the  micro- 
scope consisted  of  large  pale  vacuolated  cells  in  columns 
or  strands  not  connected  with  bile  ducts.  The  last  feature 
speaks  in  favor  of  the  diagnosis  of  adenoma.  The  damage 
to  the  organ  was  probably  considerable  and  the  portal 
circulation  must  have  been  impeded  since  passive  con- 
gestion and  ascites  were  present.  Enteritis  and  nephritis 
seemed  the  causes  of  death. 

Adenomata  or  fibroadenomata  of  bile  duct  origin  were 
seen  in  four  animals,  a  Red  Fox  (Canis  vulpes  pemi- 
sylvanicus),  a  Gray  Fox  {Canis  ciner  eo-ar  gent  ens) ,  a 
Jaguar  (Felis  onca)  and  a  Common  Deer  (Masama  vir- 
giniana).  The  first  two  present  similar  pictures,  pin- 
point to  8.  mm.,  gray,  well  outlined  areas  some  of  which 
are  clearly  cystic,  others  opaque  and  more  solid.  In  the 
first  fox  a  larger  mass  was  found  near  the  hilum.  Careful 
study  and  consultation  has  failed  to  discover  parasites  in 
these  cases,  although  their  presence  was  strongly  sus- 
pected, so  that  we  were  forced  to  conclude,  in  view  of  the 
rather  typical  microscopic  picture,  that  they  are  adeno- 
mata of  bile  duct  origin.  Their  scattered  distribution, 
but  with  a  tendency  to  be  more  numerous  beneath  the 
capsule,  corresponds  with  a  human  case  just  brought  to 
my  notice.  The  mass  in  the  liver  of  the  deer  was  single 
and  resembled  an  infarct,  with  cysts  exposed  by  cross 
section.  This  tumor  was  found  on  the  diaphragmatic 
surface  of  the  right  lobe. 


242   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Tumors  of  an  atypical,  therefore  malignant,  epithelial 
variety  were  found  four  times,  in  an  Alpaca  {Lama  pacos) 
and  three  parrakeets ;  these  birds  are  very  prone  to  have 
all  kinds  of  tumors.  The  records  of  the  first  animal  could 
not  be  as  satisfactory  as  might  be  desired  because  of  an 
advanced  state  of  decomposition  but  there  was  a  car- 
cinoma-like growth  of  the  gall-bladder  area  and  a  large 
hard  alveolated  tumor  occupying  one-half  of  the  liver. 
The  colon  had  been  involved  by  the  former,  with  perfora- 
tion. Two  of  the  parrakeets  showed  a  simple  carcinoma 
with  well  developed  fibrous  tissue  bands  running  in  all 
directions  through  the  large  mass.  The  whole  growth 
was  comparable  to  the  usual  picture  of  these  massive 
tumors  when  they  are  primary  in  the  liver.  All  these 
three  cancers  seem  to  take  their  origin  in  the  liver  cells 
but  the  third  had  such  an  interesting  involvement  of  the 
connective  tissue  that  its  minute  anatomy  will  be  given; 
it  was  denominated  adenocarcinoma  sarcomatodes. 

Undulated  Grass  Parrakeet  6  {Melopsittacus  unduJatus).  Section 
of  liver  shows  organic  capsule  normal.  Nothing  remains  of  the  original 
structure  by  which  it  might  be  recognized,  suggestion  in  places  of 
granular  cells  resembling  liver  cells  being  only  occasional  occurrences 
and  in  small  numbers.  Where  liver  cells  do  occur  they  are  highly 
granular  in  various  degrees  of  atrophy  and  show  various  grades  of 
nuclear  retrogression.  Greatest  part  of  section  consists  of  dense,  white 
fibrous  tissue  in  which  lymphocytes  are  rather  diffusely  placed  together 
with  large  numbers  of  epithelium-lined  spaces.  These  spaces  are  often 
elongated  after  manner  of  imperfect  ducts  but  are  of  irregular  form, 
have  single  layer  of  low  cuboidal  epithelium  and  richly  staining  nuclei. 
Upon  search  certain  acini  are  found  to  have  especially  hyperchromatic 
nuclei  and  penetration  of  basement  membrane.  In  such  localities  col- 
lections of  epithelial  cells  are  to  be  seen  in  plug  form  in  lymphatics 
and  acini  of  imperfect  development  of  lumen  are  found.  In  addition 
to  these  epithelial  lesions  connective  tissue  ones  are  seen,  occurring 
generally  in  restricted  localities.  The  interstitial  framework  is  seen  to 
consist  of  closely  placed  spindle  cells,  some  of  which  are  especially 
elongated  after  manner  of  imperfect  ducts  but  are  of  irregular  form, 
directed  in  a  definite,  purposeful  manner,  but  interlace  in  the  whoi'ling 
manner  noted  in  fibromas.  Nuclei  are,  however,  entirely  too  chromatic 
for  a  connective  tissue  tumor.  Whenever  a  vessel  occurs  in  these  re- 
gions its  lining  endothelium  is  always  swollen  and  nuclei  in  its  wall 


THE  LIVER  243 

will  be  proliferated  and  of  embryonic  type.  This  latter  condition  is 
apt  to  occur  in  patchy  manner,  part  of  wall  appearing  normal  and 
other  parts   containing  these   peripherated   elongated   nuclei. 

Secondary  tumors  were  observed  in  the  liver  seven 
times  as  follows:  Red  Kangaroo  {Macropus  rufus)  from 
malignant  papilloma  of  the  stomach;  Spotted  tailed 
Dasyure  {Dasyurus  ynaculatus)  from  cancer  in  the  small 
intestine;  Dorcas  Goat  {Capra  Jiircus)  from  sarcoma  in 
lymph  nodes  in  mediastinmn;  Raccoon-like  Dog  {Cams 
procyonoides)  from  mixed  tumor  of  thyroid;  Undulated 
Grass  Parrakeet  {Melopsittacus  undulatus)  from  a  brain 
tumor  probably  glioma ;  another  of  same  species  from  a 
sarcoma  of  pectoral  muscle ;  European  Robin  (Erithacus 
ruheculus)  adenoma  of  adrenal  {hypernephroma) .   . 


SECTION  VII 

THE  ALIMENTARY  TRACT,  PART  3. 
THE  PANCREAS 

The  pancreas,  an  organ  functionating  as  a  gland  with 
an  internal  secretion  and  by  pouring  a  digestive  juice  into 
the  duodenum,  remains  a  structure  of  constant  anatomy 
throughout  the  zoological  classes  under  discussion  in  that 
it  is  composed  of  compound  racemose  lobules  whose 
outlets  join  to  form  large  discharging  ducts,  and  of 
interstitial  bodies,  the  islands  of  Langerhans,  without 
connection  with  the  secreting  acini  but  having  some  rela- 
tion with  the  blood  and  lymph  vessels.  The  organ 
originates  embryologically  by  sprouts  from  the  side  of 
the  primitive  gut  just  below  the  part  destined  to  be 
stomach,  and  from  an  outbudding  of  the  common  biliary 
duct.  These  two  sprouts  or  pouches  combine  to  form  one 
organ,  but  this  does  not  necessarily  effect  a  union  between 
their  lumina.  In  some  birds  and  mammals  (Accipitres 
and  some  Ungulata)  the  lobes  of  the  pancreas  remain 
distinct  during  life,  and  the  discharging  tubules  seem  to 
empty  only  their  respective  lobes.  However,  there  is  no 
uniformity  in  the  matter,  and  indeed  the  anatomy  of  the 
ducts  is  subject  to  very  great  variation  despite  the  rather 
similar  beginnings  of  the  organ.  Those  who  are  inter- 
ested in  this  point  may  consult  Beddard,(l)  Letulle  and 
Nathan-Larrier,(2)  and  Opie  (3) ;  there  will  be  given  in 
the  following  pages  the  average  findings  of  anatomy  of 
the  gland  body  and  of  its  ducts. 

The  region  of  the  pancreas  in  lower  animals,  espe- 
cially those  which  travel  constantly  on  four  feet,  is  one  of 
great  activity,  and  the  organs  are  more  freely  movable 
than  in  the  human  being.    The  only  exception  to  the  latter 

(1)  Proc.  Zool.  8oc.  London,  1905. 

(2)  Bull.  Soc.  Anat.,  1898,  73,  491 

(3)  Amer.  Med.,  1903,  996 
244 


THE  PANCREAS  245 

part  of  this  statement  may  possibly  be  found  in  the  cats 
and  dogs,  in  which  there  are  firmer  attachments  of  the 
duodenum  and  pancreas  to  the  vertebral  column  and  the 
liver;  this  is  brought  about  by  the  short  gastrohepatic 
omentum  and  the  abrupt  curvature  of  the  duodenum 
toward  the  back,  under  the  mesenteric  stalk.  In  the 
Ungulata  and  Marsupialia  and  in  some  Rodentia,  the 
pyloric,  duodenal,  and  pancreatic  attachments  are  rela- 
tively loose,  and  torsion  of  the  pylorus  seems  to  be 
allowed  for,  since  in  these  animals  great  distention  of  the 
stomach  is  the  rule.  Among  the  Aves  the  anatomy  is 
wholly  different.  The  birds  have  no  attachment  of  the 
duodenum  and  pancreas  to  the  posterior  abdominal  wall, 
except  indirectly  through  a  narrow  strip  comparable  to 
the  gastrohepatic  omentum,  one  division  of  which  passes 
to  the  beginning  of  the  duodenum,  the  other  to  its  end,  and 
by  a  thin  tail  of  pancreas  which  goes  toward  the  spleen. 
The  bulk  of  the  pancreas  lies  in  the  U  made  by  the  long 
free  duodenal  loop,  the  two  organs  being  covered  by  the 
serous  membranes  forming  the  middle  abdominal  sac.  It 
will  be  seen  from  the  foregoing  that  the  movability  of  the 
pancreas  is  considerable — a  highly  necessary  provision, 
because  the  stomach  and  duodenum  are  also  movable  and 
subject  to  distention  by  food  and  alteration  of  position 
during  flight. 

In  the  class  Mammalia  there  are  usually  two  ducts, 
one  entering  the  duodenum  in  combination  with  the  bile 
duct,  the  other  variously  above  or  below  this  common 
opening.  As  will  be  seen  in  Table  14,  however,  there  are 
several  exceptions  to  this  statement,  there  being  but  one 
duct  opening  independently  of  the  bile  duct.  The  general 
anatomy  is  closely  similar  throughout  this  class,  so  I  shall 
confine  my  notes  to  the  exceptions  from  the  general  rule, 
especially  where  they  seem  to  be  of  importance  in  the 
etiology  of  pancreatic  lesions. 

In  the  class  Aves  the  pancreas  consists  usually  of  two 
or  three  distinct  lobes  lying  one  in  front  and  two  behind 


246   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

the  cleft  between  the  limbs  of  the  duodenal  loop,  and  it 
discharges  its  secretion  into  the  duodenum  by  two  or 
three  ducts  separately,  and  almost  invariably  above  the 
bile  duct  openings.  One  duct  always  opens  near  the  top 
of  the  distal  end  of  the  duodenal  loop,  near  the  bile  duct. 
In  the  gallinaceous  birds  that  have  a  bile  duct  opening 
into  the  duodenum  near  the  pylorus,  there  is  usually  a 
pancreatic  duct  opening  there  also.  In  some  birds  a  third 
duct  passes  from  the  body  of  the  pancreas  to  the  duo- 
denum at  different  places  along  the  loop.  It  does  not 
seem  probable  that  dislocation  of  the  duodenal  loop  would 
seriously  interfere  with  the  passage  of  the  pancreatic 
secretions,  since  the  gland  is  so  intimately  related  with 
the  duodenal  serosa,  but  obstruction  to  the  biliary  flow 
due  to  changes  in  position  of  the  intestine  is  easier 
because  the  bile  duct  is  separate  and  loose  and  arises 
from  the  end  of  the  gall-bladder.  The  ducts  of  both  these 
structures  pass  very  obliquely  through  the  duodenal  wall 
a  matter  of  importance,  as  will  be  seen  when  discussing 
the  infiltrative  forms  of  enteritis.  The  gall-bladder  is 
not  present  in  all  birds,  but  this  is  probably  of  no 
importance,  as  the  hepatic  ducts  are  wide  and  run 
directly  from  the  liver  to  the  duodenum.  The  pancreatic 
ducts  are  short  and  are  closely  bound  around  by  glandu- 
lar tissue  up  to  a  place  quite  close  to  their  entrance  into 
the  intestine. 

The  musculature  of  the  gall-bladder  and  the  ducts 
seems  comparable  in  mammals  and  birds,  and  a  con- 
strictor or  sphincter  usually  called  the  muscle  of  Oddi, 
is  present  in  all  but  pigeons  (Oddi).  There  may  be  found 
also  muscular  fibres  in  the  major  ducts  of  the  pancreas, 
but  they  are  not  so  heavy  nor  distributed  so  definitely  as 
similar  tissue  in  the  bile  duct  walls.  The  mucosa  of 
the  pancreatic  duct  is  much  more  folded  in  birds  than  in 
mammals,  seemingly,  therefore,  more  adapted  to  obstruc- 
tion by  swelling  from  any  cause. 


THE  PANCREAS  247 

Passerine  birds  have  two  pancreatic  ducts  usually  on 
the  ascending  loop  of  the  duodenum,  or  there  may  be  one 
ahead  of  the  pyloric  biliary  duct.  The  picarian  varieties 
possess  three  ducts  as  a  rule,  one  near  the  beginning  of 
the  pylorus,  one  near  its  end  and  a  third  of  inconstant 
location.  Owls  have  a  system  like  Passeres,  but  the  rela- 
tion between  the  organ  and  the  intestinal  loop  is  looser 
and  the  ducts  are  wider.  ColumbsB  have  two  pancreatic 
ducts  in  the  ascending  limb  of  the  duodenum.  Gallinaceous 
varieties  have  a  double  biliopancreatic  system,  a  duct  of 
each  kind  entering  the  descending  and  the  ascending 
duodenal  reaches,  with  the  biliary  placed  after  the  pan- 
creatic in  each  instance.  Accipitres  have  always  two 
and  oftentimes  three  ducts  as  do  Anseres,  both  orders 
frequently  having  the  third  duct  opening  at  the  bottom 
of  the  duodenal  loop  where  stagnation  can  and  does  occur. 
FulicariaB  have  usually  three  ducts. 

The  foregoing  are  the  orders  presenting  pancreatitis 
and  therefore  those  whose  anatomy  concerns  this  study 
directly.  The  irregularity  in  number  and  arrangement  of 
ducts  continues  through  all  the  avian  orders  which  show 
a  greater  aberration  from  standards  than  do  the  mam- 
mals. Theoretically  the  birds  should  cast  some  light  upon 
the  unsettled  question  of  the  causes  of  pancreatitis,  and 
as  a  matter  of  fact  such  a  result  seems  to  have  been 
realized.  In  1915  I  published  an  article  upon  a  study  of 
this  subject  which  indicated  that  acute  inflammations  of 
this  organ  may  arise  via  the  lumen  of  the  duodenum  and 
pancreatic  ducts,  while  chronic  processes  were  the  result 
of  periductal  passage  of  pathogenic  agents.  Further 
study  would  seem  to  indicate  that  disease  of  the  biliary 
tract  is  of  importance  in  lesions  of  the  pancreas  since  a 
decidedly  large  number  of  cases  is  found  in  mammals, 
where  the  relation  of  ducts  is  definitely  more  intimate 
than  in  birds.     The  work  of  Archibald, (4)  Deaver  and 

(4)   Surg.  Gyn.  and  Obst.,  1919,  28,  p.  529. 


248  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Sweet, (5)  and  Judd  (6)  seem  to  agree  with  the  findings 
upon  our  material.  This  need  not  be,  however,  in  discord 
with  the  idea  that  acute  inflammation  is  superficial  in 
origin,  chronic  lesions  deep  or  lymphogenic.  The  discus- 
sion will  be  resumed  in  a  subsequent  paragraph. 

The  amount  of  pancreas  to  be  found  in  birds  is  greater 
than  that  in  manmials.  According  to  our  figures  the  organ 
represents  ^/4ooth  of  the  body  weight  in  the  former  and 
Voooth  in  the  latter.  These  figures  are  averages  of  a  small 
number  of  instances  and  are  not  final.  It  is,  however, 
obvious  to  casual  daily  observation  that  birds  as  a  class 
have  a  large  pancreas. 

The  minute  structure  of  the  organ  is  governed  by  the 
same  general  rules  throughout  the  two  classes  under  con- 
sideration. Birds  do  not  have  as  many  interstitial  islands 
as  do  mammals,  but  they  are  more  compact  and  seem 
more  definitely  constructed  of  coiled  tubules.  In  so  far 
as  the  internal  structure  of  the  organ  is  concerned  there 
has  not  developed  in  our  study  pathology  peculiar  to  any 
animal.  The  importance  of  the  ducts  and  position  of  the 
organ  will  be  discussed  later. 

Recognition  of  pancreatic  disease  during  life  is  prac- 
tically impossible.  In  human  medicine  the  signs  and 
symptoms  are  vague  and  inconstant, (7)  diagnosis  often 
being  a  matter  of  exclusion.  Veterinarians,  except  under 
the  best  hospital  conditions  make  no  attempt  to  diagnose 
pancreatic  lesions  but,  since  the  improvement  of  surgical 
practice,  at  times  operate  upon  cases  of  evident  pain  and 
distention  which  prove  to  be  pancreatitis.  These  things 
were  evident  in  a  deer  that  I  saw  and  that  died  on  the 
following  day  from  acute  hemorrhagic  pancreatitis;  I 
made  no  attempt  at  this  diagnosis,  believing  it  to  be  acute 
tympanites.  The  feces  were  normal,  according  to  the 
judgment  of  persons  qualified  to  give  an  opinion. 

(5)  Jour.  A.M. A.,  1921,  77,  194.  ~        ~~ 

(6)  Ibid.,  197. 

(7)  Garrod,  Schorstein  Led.,  1920. 


THE   PANCREAS  249 

The  condition  of  the  pancreas  at  autopsy  on  animals 
not  dying  with  lesions  of  this  organ  deserves  some  atten- 
tion since  it  may  confuse  the  uninitiated.  If  the  organ  be 
seen  in  its  normal  resting  stage  shortly  after  death,  it  is 
not  difficult  to  recognize  the  condition  as  normal  for  the 
species.  Activity  is  indicated  by  a  darker  or  redder  color 
and  an  increase  of  consistency.  In  carnivorous  or 
omnivorous  animals  and  birds  the  pancreas  in  this  state 
is  a  body  with  a  distinct  bulky  character,  whereas  in 
strictly  herbivorous  varieties,  especially  ungulates,  the 
structure  is  diffusely  pink  and  doughy.  This  is  impor- 
tant since  the  early  stages  of  self-digestion  and  decompo- 
sition assume  this  same  character  in  all  varieties,  while 
later  stages  present  a  deep  red,  swollen,  wet  organ.  These 
appearances  must  be  differentiated  from  acute  hemor- 
rhages or  inflammations,  a  distinction  based  upon  actual 
local  blood  collections  or  extravasations  and  areas  of 
degeneration  in  true  disease.  Oftentimes  differentiation 
must  be  made  under  the  microscope  and  in  advanced 
decomposition,  determination  is  impossible.  When  there 
is  torsion  of  the  stomach,  notably  in  ungulates,  the  pan- 
creas is  often  found  decidedly  congested.  This,  it  seems, 
is  due  to  a  twist  of  the  duodenum  and  passive  congestion 
of  it  and  the  pancreas — the  only  simple  explanation 
despite  the  apparent  provision  for  a  high  degree  of 
mobility,  as  already  explained.  The  organ  is  nearly 
always  mildly  congested  in  severe  grades  of  acute  duo-, 
denitis,  although  it  need  not  be  pathologically  involved. 
It  is,  however,  noteworthy  that  the  pancreas  is  an  organ 
with  a  low  morbidity  index,  especially  when  one  considers 
its  proximity  to  a  structure  showing  the  highest  disease 
index  in  the  body,  the  intestine.}  The  succeeding  para- 
graphs will  reveal  in  comparison  to  other  organs  only  a 
small  number  of  cases  of  degeneration,  inflammation  and 
tumors.  This  has  been  ascribed  to  the  freedom  of  blood 
supply  and  the  power  of  tryptic  digestion. 

17 


250   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

An  expression  of  this  relative  immunity  to  pathologic 
change  is  met  in  analyzing  the  data  upon  the  simplest 
lesions,  degenerations,  to  be  expected  in  many  states  of 
disease.  Only  a  small  number  of  cases  present  them- 
selves, and  they  are  under  expected  conditions,  namely 
in  association  mth  acute  general  infection,  sometimes 
definitely  septicemic  in  nature.  About  half  of  them  were 
discovered  microscopically,  affecting  the  islands  of 
Langerhans  in  vacuolization  or  granular  disintegration. 
Focal  necroses  of  the  organ  were  met  four  times, 
three  turkeys  and  a  cockatoo.  It  is  noteworthy  that 
all  these  birds  had  some  involvement  of  the  liver, 
twice  a  complete  acute  hepatitis  and  twice  a  cholangitis. 
This  is  the  more  interesting  since  we  shall  learn 
that  the  liver  is  less  often  involved  in  avian  than  in 
mammalian  pancreatitis.  Hemorrhages  occur  occasion- 
ally in  the  pancreas  in  acute  general  infections  and  are 
seen  in  acute  inflammations  of  the  intestines ;  the  percent- 
age incidence  with  the  latter  is,  however,  very  small. 
Pancreatic  apoplexy  proper  has  not  occurred,  for  all  the 
instances  of  large  hemorrhage  into  the  organ  have  been 
combined  with  changes  forcing  a  classification  of 
acute  pancreatitis. 

Panckeatitis. 

Pancreatitis  in  the  acute  form  is  divided  by  many 
writers  into  exudative,  hemorrhagic  and  necrotizing, 
while  for  the  chronic  variety  an  inter-  and  intra-acinus 
form  has  been  described.  It  is  questionable  whether  it  is 
fair  in  acute  cases  to  focus  attention  by  special  nomen- 
clature on  different  macroscopic  pictures,  unless  it  be  for 
descriptive  purposes  solely,  since  there  is  nothing  at  hand 
to  indicate  that  differing  agents  cause  one  kind  every 
time.  The  physical  findings  seem  to  depend  rather  upon 
the  speed  of  operation  of  the  causation  than  upon  its 
essence.  Sudden  obstruction  of  the  pancreatic  duct  is 
believed  to  produce  necrotizing  processes  to  which  hemor- 


THE  PANCREAS  251 

rhage  may  be  added  by  digesting  of  blood  vessels.^ 
Exudative  cases  seem  due  to  extension  of  ulcerative 
inflammation,  from  a  perforated  gastric  ulcer  for 
example,  to  whicb  digestive  pancreatitis  may  be  added. 
The  interacinus  chronic  inflammations  are  usually  con- 
sidered as  due  to  obstruction  or  infection  through  the 
biliary  or  pancreatic  ducts  whereas  vascular  disease  pro- 
duces intra-acinus  connective  tissue  overgrowth.  Analysis 
of  the  records  of  this  laboratory  would  seem  to  indicate 
that  necrotizing  and  hemorrhagic  processes  belong 
together,  exudative  in  a  class  by  themselves,  and  that 
chronic  disease  may  be  either  interlobular  or  intra-acinar 
without  regard  to  associated  pathology.  I  have  therefore 
studied  our  cases  from  this  standpoint. 

Pancreatitis  has  occurred  in  thirty-eight  mammals 
and  birds  among  the  5365  autopsies,  an  incidence  of 
0.7  per  cent. ;  class  incidence  in  mammals  twenty-seven  or 
1.5  per  cent.;  birds  eleven  or  .3  per  cent.  (Table  14.) 
Among  the  higher  class  all  the  important  orders  are  rep- 
resented, but  by  no  means  in  equal  degree,  whereas  in  the 
birds,  less  than  half  of  the  orders  are  listed,  with  the 
important  Psittaci  missing,  despite  a  high  death  rate. 

It  is  perhaps  well  to  be  guarded  in  stating  the  relative 
vulnerability  of  the  pancreas  in  various  orders,  but  one 
cannot  avoid  the  observation  that  Carnivora  stand  well 
in  advance  of  the  others  (3.  per  cent,  of  autopsies),  to  be 
followed  by  Ungulata  (1.9  per  cent.)  and  Rodentia 
(1.7  per  cent.).  Nor  can  one  fail  to  see  that  mammals 
have  inflammations  of  this  organ  five  times  as  often  as 
do  birds. 

Further  analysis  of  the  data  leads  into  a  consideration 
of  the  anatomy  of  the  viscus  in  terms  of  the  acceptable 
theories  of  the  origin  of  the  lesion.  It  is  commonly 
believed  that  infection  of  the  gland  occurs  by  passage  of 
organisms  through  the  duct  opening  in  the  intestines, 
especially  when  there  is  swelling  of  the  mucosa  of  both. 
For  the  human  being  the  idea  is  current  that  infection  or 


252  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 


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THE  PANCREAS 


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254  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 


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THE  PANCREAS  255 

obstruction  of  the  common  bile  duct  may  spread  to  the 
pancreatic  duct,  and  that  mechanical  or  inflammatory 
obstruction  of  the  papilla  of  Vater  may  permit  the  bile  to 
pass  up  the  pancreatic  duct.  This  theory  is  based  upon 
certain  observations, notably  those  of  Opie,  in  cases,  where 
a  gall  stone  obstructed  the  ampulla,  bile  entered  the 
pancreas  and  acute  pancreatitis  arose,  partly  by  the  acti- 
vating action  of  the  bile  upon  the  pancreatic  juice  and 
partly  by  bacteria  introduced  at  the  same  time.  This 
method  of  origin  is  perhaps  accepted  in  most  quarters, 
but  there  are  some  who  believe  that  infection  of  the  gland 
may  occur  by  the  infiltration  of  lymphatics  around  the 
pancreas  by  disease  of  adjacent  parts — gall-bladder, 
stomach  or  lymph  nodes.  Quotation  has  already  been 
given  to  reference  literature,  and  I  shall  not  go  further 
into  theory  except  in  pointing  out  how  our  material  may 
help  to  answer  the  question. 

In  the  first  place  it  seems  perfectly  obvious  that  infec- 
tion might  in  any  animal  travel  from  the  intestine  to  the 
pancreas  via  its  ducts,  the  main  question  to  be  settled 
being  the  relative  importance  of  the  infection  via  the  bile 
duct.  Let  us  now  see  if  the  variations  in  anatomy  will 
cast  any  light  upon  the  matter. 

Among  the  1860  mammals,  there  are  as  far  as  I  can 
determine  1275  which  have  one  pancreatic  duct  opening 
in  conjunction  with  the  bile  duct,  585  in  which  the  former 
has  an  intestinal  opening  independent  of  the  latter. 
Among  this  1275  there  are  nineteen  cases  of  pancreatitis, 
while  among  the  585  there  are  eight  cases,  or  as  14.9  to 
13.6.  All  Aves  have  separate  biliaiy  and  pancreatic  ducts 
and  relatively  little  pancreatitis,  although  frequently 
suffering  with  its  most  common  accompaniment,  namely 
duodenitis.  There  is  therefore  some  evidence  that  more 
pancreatitis  occurs  when  there  is  a  physical  proximity 
or  combination  of  bile  and  pancreatic  ducts. 

Active  infections  of  the  biliary  system  in  relation  to 
pancreatitis  are,  however,  not  as  conspicuous  as  might  be 


256   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

expected.  In  the  maniinals  twelve  of  the  twenty-seven 
cases  showed  cholecystitis  or  cholangitis;  in  every 
instance  the  form  of  pancreatitis  was  acute.  Among  the 
eleven  avian  cases  four  showed  inflammation  of  the 
biliary  channels,  but  not  of  the  bladder.  Hepatic  cirrhosis 
was  observed  four  times.  In  a  thrush  and  a  skunk 
obvious  infectious  cirrhosis  existed,  and  in  both  a  necro- 
tizing pancreatitis  was  found.  A  badger  suffered  with 
atrophic  cirrhosis  of  the  liver  and  a  chronic  pancreatitis 
wdtli  acute  exacerbation.  A  deer  showed  marked  peri- 
lobular fibrosis  wT.th  a  recent  hemorrhagic  pancreatitis 
probably  due  to  duodenal  torsion.  Nothing  very  distinc- 
tive is  to  be  found  in  these  cases,  but  they  merely  make  the 
total  of  involvements  of  the  liver  and  its  adnexa  up  to 
tw^enty.  It  is  to  be  emphasized  that  pancreatitis  was  not 
associated  with  lithiasis  in  ducts  or  bladder  as  described 
on  page  240.  Peripheral  cholecystitis  and  plastic  inflam- 
mations about  the  pylorus  and  lesser  omentum  are 
exceedingly  rare  in  wild  animals,  w^hile  they  are  not  com- 
mon in  human  surgical  practice.  They  did  not  occur  at 
all  in  mammals  in  this  series,  the  only  external  inflamma- 
tions being  in  lymph  nodes  in  cases  of  frank  infectious 
character.  There  were  distinct  adhesions  between  liver, 
duodenum  and  pancreas  in  two  birds,  one  with  acute,  the 
other  with  chronic  pancreatitis. 

In  so  far  as  the  kind  of  pancreatitis  is  concerned  mam- 
mals had  twenty-two  acute  and  six  chronic  forms,  one 
animal  having  the  former  implanted  on  the  latter,  w^hile 
birds  had  five  acute  and  six  chronic.  The  preponderance 
of  acute  over  chronic  lesions  in  mammals  again  recalls 
the  association  of  the  biliary  and  pancreatic  ducts,  but 
if  one  expect  that  such  a  relation  establishes  acute  inflam- 
mation, the  relatively  high  figures  for  Rodentia  and 
Ungulata,  with  a  single  duct  removed  from  the  bile  duct 
conflict  with  the  data  for  orders  having  two  ducts  such 
as  the  Carnivora.  Every  case  in  the  former  orders  was 
of  acute  nature ;  only  two  had  any  hepatic  disease,  four 


THE  PANCREAS  257 

had  lymph  gland  hyperplasia  in  the  pancreatic  region, 
and  seven  had  acute  enteritis.  The  preponderance  of 
acute  over  the  chronic  cases  in  mammals  and  the  nearly 
equal  number  in  birds  is,  however,  apparent. 

The  collateral  pathology  with  the  most  definite  rela- 
tionship to  pancreatitis  is  enteritis  and  one  may  say  that 
the  former  occurs  in  proportion  to  the  incidence  of  the 
latter.  Acute  forms,  twenty-seven,  were  associated  with 
acute  enteritis  nineteen  times.  Chronic  enteritis  was 
found  with  chronic  pancreatitis  in  six  of  eleven  cases. 

There  is  a  rough  relationship  between  the  type  of 
pancreas  and  the  nature  of  the  lesions.  The  organ  may 
be  divided  for  this  purpose  into  the  compact  organ  firmly 
held  in  place  by  attachments  to  the  lesser  omentum, 
spleen  and  duodenum,  and  the  velamentous  organ  which 
spreads  a  considerable  distance  along  the  duodenum  and 
sends  out  digitations  into  the  mesentery  and  thin 
processes  toward  the  spleen.  The  first  type  is  seen  in 
primates  and  carnivores  and  birds,  while  the  second 
is  characteristic  of  rodents,  ungulates  and  marsupials. 
The  compact  variety  showed  all  but  one  of  the  chronic 
cases  while  the  loose  organ  was  atfected  by  the  acute  pan- 
creatitis in  twelve  of  thirteen  cases. 

The  microanatomy  of  the  cases  may  throw  a  little 
light  upon  our  subject.  T  was  able  to  see  the  duct  in  one 
case  of  acute  pancreatitis  (bear).  It  showed  a  simple 
catarrhal  inflammation  with  a  very  moderate  circumfer- 
ential round  cell  increase.  The  destruction  of  glandular 
areas  by  edema,  hemorrhage  and  necrosis  offers  nothing 
of  importance  except  in  a  few  birds.  In  these  the  necrosis 
is  more  definite  about  cross  sections  of  ducts,  and  the 
islands  of  Langerhans  are  frequently  spared  until 
necrosis  is  locally  complete.  In  a  case  of  chronic  pancre- 
atitis in  a  bird,  a  cross  section  of  pancreatic  duct  was 
found  in  the  intestinal  wall ;  a  chronic  catarrhal  and  infil- 
trative enteritis  existed  in  this  specimen.  A  very  definite 
mantle  of  round  cells  was  found  about  the  duct  while  the 


258  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

mucosa  showed  no  change,  although  the  lumen  seemed 
large.  Interlobular  fibrosis  was  the  rule,  only  one  case 
of  intra-acinar  pancreatitis  being  encountered ;  this  speci- 
men, a  bird,  showed  great  distortion  of  the  acini  and  of 
the  islets. 

The  study  of  comparative  pathology  of  pancreatitis 
does  not  settle  its  etiology,  but  some  very  suggestive 
facts  may  be  learned.  The  association  of  hepatic  and 
biliary  disease  and  of  enteritis  in  the  causation  of  pan- 
creatitis seems  amply  confirmed,  and  the  latter  factor  is 
in  our  series  numerically  the  greater.  It  is  suggestively 
showTi  that  inflammations  of  the  pancreas  occur  more 
frequently  in  the  zoological  class  in  which  the  ducts  of  the 
organ  and  of  the  liver  empty  into  the  duodenum  together 
or  in  close  association.  Morover,  infections  of  the  liver 
and  adnexa  are  very  important  in  the  mammals,  more  so 
than  birds  in  which  enteritis,  notably  chronic  in  type, 
usually  accompanied  the  involvement  of  the  pancreas. 
This  is  consistent  with  the  incidence  of  bile  tract  disease 
as  already  discussed  under  that  subject,  and  it  is  inter- 
esting to  note  that  the  mammals  showing  the  greatest 
number  of  cases  of  choledochitis  and  cholangitis  also 
show  the  high  case  incidence  of  pancreatitis.  The  birds 
that  have  bile  tract  disease  have  little  pancreatic  disease. 
These  facts  when  considered  in  connection  with  the  free 
biliary  supply  of  the  avian  duodenum,  the  disassociation 
of  the  ducts  of  the  two  glands  and  the  close  apposition  of 
the  pancreas  to  the  duodenal  wall,  suggest  strongly  that 
direct  infection  of  the  pancreas  can  occur  from  the 
intestinal  wall  along  the  walls  of  the  ducts  perhaps  via 
the  lymphatics.  This  is  supported  by  the  observation  of 
at  least  one  case  in  which  there  was  a  definite  inflamma- 
tion under  the  adventitia  of  the  pancreatic  duct,  its 
mucosa  being  normal.  The  study  also  suggests  that  acute 
pancreatitis  is  more  often  associated  w^th  acute  lesions 
in  the  intestines  and  with  hepatic  or  gall-bladder  disease, 
and  that  chronic  pancreatitis  seems  more  often  the  result 


Kk,.   20. — ADENOMA  OK  PANCRKATIC    DUCTS.      CORSAC   FOX   (CANIS  CORSAC). 


THE  PANCREAS  259 

of  chronic  or  repeated  intestinal  inflammation.  Periph- 
eral inflammation  such  as  occurs  in  gastric  or  duodenal 
ulcers,  has  not  been  encountered  in  a  distinct  character  so 
that  its  value  cannot  be  estimated. 

Cystic  change  in  the  pancreas  has  been  observed  a  few 
times,  but  never  a  large  visceral  collection  or  the  so-called 
extra-pancreatic  cysts  of  the  omentum.  One  acinus  cyst 
was  seen  in  a  drake,  one  congenital  cyst  in  a  lark,  and 
multiple  ductal  cysts  were  seen  in  a  baboon  and  a  duck. 
The  parenchyma  in  all  cases  seemed  entirely  capable 
of  functionating.  \ 

Tumors. 

Tumors  of  the  pancreas  have  been  three  in  number, 
two  being  of  academic  interest  only.  One  of  these  con- 
cerned an  apparent  adenoma  of  the  ducts  within  the 
organ,  discovered  microscopically  in  the  sections  from  a 
Corsac  Fox  (Canis  cor  sac).  This  is  the  only  specimen 
we  have  had,  and  I  can  find  no  description  of  the  normal 
microanatomy  of  this  species,  so  that  with  a  knowledge 
that  certain  carnivores  have  convoluted  ducts,  the 
determination  is  made  tentatively ;  it  corresponds  micro- 
scopically to  a  ductal  adenoma.  The  pancreas  of  a 
raccoon  {Procyon  lotor)  showed  a  true  adenoma  of 
glandular  acini  as  two  separate  but  closely  applied  firm 
nodules  surrounded  by  a  capsule,  and  with  distorted  acini 
as  seen  under  magnification.  The  most  important  tumor 
was  found  in  an  Indian  Paradoxure  {Paradoxurus  niger), 
an  adenocarcinoma  involving  the  head  of  the  organ, 
enlarging  it  to  twice  its  normal  size ;  there  were  no  metas- 
tases. The  animal  suffered  also  with  an  infective  hepatic 
cirrhosis  of  recent  origin,  chronic  nephritis  of  the  arterio- 
sclerotic type,  chronic  fibroid  splenitis.  No  obstruction 
to  the  biliary  channels  existed. 


SECTION  VII 

THE  ALIMENTARY  TRACT,  PART  4. 
THE  PERITONEUM 

This  visceral  envelope  is  principally  important 
because  of  the  fatal  character  of  its  acute  inflammations. 
In  man  peritonitis  of  acute  origin  and  type  is  commonly 
secondary  to  a  focus  of  progressive  inflammation  in  some 
abdominal  organ  and  usually  speaks  for  the  \^rulence  of 
the  primary  disease  and  for  the  low  resistance  of  the 
serous  membrane.  Because  of  this  vulnerability,  greater 
foresight  is  attempted  to  prevent  the  extension  of  acute 
intra-abdominal  inflammations  and  under  operative  con- 
ditions punctilious  care  is  used  to  avoid  contamination  of 
the  general  peritoneal  cavity.  Involvement  of  the  peri- 
toneum in  septicemic  states  is  relatively  uncommon  in 
man,  but  seemingly  more  frequent  in  the  lower  mammal. 
This  surface  seems  more  resistant  to  infection  at  oper- 
ation in  the  lower  animals  since  post-operative  peritonitis 
after  castration  and  experimental  procedure  is  certainly 
infrequent ;  our  data  will  permit  no  percentage  figures  of 
vulnerability  under  such  conditions.  Judging,  however, 
from  the  number  of  times  at  which  the  diagnosis  of  acute 
peritonitis  has  been  made,  the  lower  animal  has  a 
decidedly  low  percentage  resistance  although  its  patho- 
logical states  are  primary  or  secondary  to  conditions 
unusual  in  man.  Among  the  5365  autopsies  acute  peri- 
tonitis appears  in  the  diagnoses  137  times  or  2.4  per  cent. ; 
mammals,  57  or  3  per  cent. ;  birds,  80  or  2.3  per  cent. ; 

The  exact  causes  are  usually  obvious,  practically 
always  so  in  human  medicine,  but  a  number  of  cases 
escape  adequate  explanation.  From  a  practical  stand- 
point two  origins  are  important  to  zoological  collections, 

260 


THE  PERITONEUM  261 

trauma  and  intestinal  perforations  by  sharp  objects.) 
Ungulata  frequently  suffer  abdominal  injuries  in  fighting,  1 
as  do  rodents.  Peritonitis  sometimes  supervenes  even  in 
the  absence  of  penetrating  wounds,  probably  by  reason  of 
damage  to  the  intestine  whereby  its  permeability  is 
increased.  Pointed  objects  are  frequently  swallowed  by 
animals  and  perforation  occurs.  The  danger  of  feeding 
split  bone  to  carnivores  is  well  known ;  some  years  ago 
we  lost  two  tigers  and  a  lion  in  this  manner. 

Ileus,  in  one  of  the  several  foniis,  has  been  an 
occasional  cause  of  peritonitis  in  primates  and  ungu- 
lates. The  extension  of  purulent  inflammation,  abscesses 
and  the  like  is  easy  to  understand,  but  we  have  seen 
several  cases  of  apparent  extension  from  enteritis  with- 
out perforation.  The  reason  for  this  is  probably  in  the 
kind  of  enteritis.  Monkeys  with  amoebiasis  and  gallina-— 
ceous  birds  with  enterohepatitis  have  supplied  most  of 
the  cases,  these  infestations  of  the  gut  wall  being  deep 
and  spreading  so  that  a  chance  is  afforded  to  penetrate 
the  serosa  along  blood  and  lymph  vessels.  One  case  in  a 
deer  seemed  to  originate  from  a  simple  catarrhal  colitis ; 
trichocephalus  in  the  colon  may  have  helped.  Parasites 
are  not  very  potent  in  causing  an  acute  peritonitis,  but 
aggravate  the  action  of  other  agents.  Septicemic  states 
are  at  the  bottom  of  24  per  cent,  of  our  cases  of  peri- 
tonitis. This  is  particularly  true  of  birds,  it  being 
recognized  that  their  acute  general  infections  frequently 
have  such  an  effect,  but  the  primates  and  carnivores  also 
have  a  vulnerable  peritoneum  when  septicemia  exists. 
The  principal  outstanding  visceral  lesions  in  the  mam-^ 
malian  cases  is  pneumonia;  .in  birds  it  is  cholera  andj 
plague.  The  rupture  of  eggs  in  birds  lays  the  foundation! 
of  a  peritonitis,  while  bacteria  from  the  oviduct  or  cloaca 
complete  the  process.  Chronic  peritonitis  is  not  common. 
It  is  usually  due  to  parasites  or  to  tuberculosis.    There  / 


262  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

has  been  observed,  however,  no  complete  general 
involvement  of  the  peritoneal  cavity  including  the  liver 
and  spleen,  sometimes  called  ' '  sugar-icing, ' '  and  believed 
to  be  tuberculous  in  origin.  The  only  tumor  found  in  very 
close  association  with  the  peritoneum  occurred  in  a 
Chapman's  Zebra  {Equus  hurchelli  cliapmani)  in  which 
animal  a  iibromyoma  seemed  to  spring  from  a  loop  of 
intestine  and  grow  away  from  the  gut  wall.  It  was  under- 
going myxoid  change. 


SECTION  VIII 
THE  URINAEY  TRACT 

The  Kidney. 

The  kidneys,  ureters,  bladder  and  urethra  remain  com- 
parable in  all  mammals  excepting  the  monotremes  where 
there  is  no  urinary  passage  through  genital  openings, 
the  urine  being  ejected  through  the  vesicoanal  pouch,  a 
sort  of  cloaca.  In  the  bird  the  first  two  parts  remain  as 
in  mammals  while  the  ureters  terminate  in  a  hernia-like 
pouch  of  the  rear  wall  of  the  cloaca.  It  would  seem 
from  this  arrangement  that  ureteral  transmission  of 
infection  from  the  anal  area  to  the  kidneys  would  be 
facilitated  in  the  lowest  mammals  and  in  the  Aves.  Varia- 
tions in  size,  shape  and  position  exist  to  a  minor  extent  in 
the  higher  orders  but  in  all  forms,  the  system  remains  a 
post-peritoneal  structure. 

Differences  in  construction  are  to  be  seen  for  example, 
in  the  single  pyramidal  kidneys  of  marsupials  and  certain 
rodents, in  the  lobulated  organ  of  bears, cattle  and  seals,  in 
the  twisted  viscus  of  horses,  but  these  gross  appearances 
do  not  destroy  the  uniform  scheme  upon  which  the  func- 
tionating unit  is  built.  The  single-lobed  kidney  discharges 
all  its  collecting  tubules  into  one  calyx  while  the  multiple 
pyramids  of  the  lobulated  organ  are  fitted  with  individual 
calices  which  in  turn  empty  into  the  pelvis  proper;  this 
is  true  whether  the  lobulations  are  retained,  as  in  the 
bear,  or  are  smoothed  out  in  the  course  of  development  as 
in  man.  The  secretory  tubule  remains  in  essentially  the 
same  form  in  all  kidneys ;  the  modern  idea  of  its  anatomy 
may  be  found  in  the  work  of  Huber(l). 

The  most  decided  example  of  the  lobulated  kidney  is 
to  be  found  in  thci  class  Aves,  wherein  the  organ  consists 

(1)  Anatomical  Record,  1917,  13,  p.  305,  On  the  morphology  of  the 
renal  tubule  in  the  vertebrates. 

263 


264   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

of  two  or  three  large  lobes  lying  in  concavities  of  the 
sacrum,  each  lobe  being  made  up  of  tiny  lobules.  The 
latter  appear  to  the  unaided  eye  as  fairly  distinct 
di\dsions  whether  viewed  on  the  exterior  or  by  cross 
section.  There  is  a  cortex  and  a  medulla  to  each,  the 
separation  being  clear  in  a  large  specimen,  vague  in  a 
small  one.  These  lobulations  are  quite  well  observed 
when  the  kidney  is  full  of  urates,  a  common  finding  in 
birds.  Magnification  of  the  ai'ian  kidney  reveals  an 
apparently  simpler  tubular  arrangement  than  is  found 
for  mammals  (2)  (3),  yet  the  relation  of  vascular  plexuses 
and  secreting  tubules  remains  similar.' 

From  the  standpoint  of  comparative  pathology  atten- 
tion can  be  drawn  to  the  glomerulus,  to  the  interstitial  tis- 
sue, and  to  the  character  of  the  epithelium.  The  tuft  of 
intricately  wound  capillaries  called  the  glomerulus  has 
always  been  viewed  as  the  part  of  the  secretory  unit 
chiefly  concerned  in  urine  production  whether  one  accept 
the  older  idea  that  it  excretes  only  fluid  or  the  modern 
belief  of  many  observers  that  all  parts  of  the  urine  go 
out  through  it.  In  the  mammal  the  tuft  is  closely  wound, 
is  surrounded  by  a  distinct  space  and  a  limiting  membrane 
of  appreciable  "width  called  Bowmian's  capsule;  all  this 
so-called  Malpighian  body  has  a  breadth  varying  from 
120  to  300  micra.  There  is  however  great  variation  in 
the  size  of  this  body  when  seen  in  the  peripheral  and 
deeper  zones  of  the  same  organ,  amounting  at  times  to 
seventy  per  cent,  of  the  diameter.  The  capillary  con- 
geries forming  the  avian  tuft  is  by  no  means  so  delicate 
and  one  can  see  individual  capillaries  Avith  more  ease.  It 
may  be  impossible  to  discover  a  space  between  the  tuft 
and  its  exceedingly  delicate  capsule,  the  latter  being 
usually  applied  closely  to  the  vascular  corpuscle.  The 
whole  breadth  varies  from  70  to  140  micra  with  an  average 

(2)  Policard,  C.  R.  Assoc.  Anat..  1910,  12,  57. 

(3)  Huber,  Anat.  Record,  1916,  10,  201. 


THE  URINARY  TRACT  265 

of  110.     There  is  more  uniformity  in  size  than  in  the 
mammalian  organ. 

Supporting  tissues  between  the  tubules  seem  less 
definitely  nuclear  in  the  bird  than  in  the  mammal,  at  least 
in  so  far  as  connective  tissue  is  concerned,  there  being  in 
the  former  only  a  few  groups  of  mononuclears  to  be  seen 
in  the  cortex.  Perivascular  tissues  are  reasonably  rich. 
The  epithelium  of  proximal  and  distal  tubules  is  not 
easily  fixed  by  our  customary  laboratory  techniques,  the 
best  results  being  obtained  by  Zenker's  fluid.  As  seen  in 
a  routine  specimen  of  a  normal  organ  it  is  vacuolated  or 
very  palely  stained.  The  individual  cells  stand  out  clearly 
and  many  present  a  pointed  end  to  the  tubular  lumen. 

Kidney  Weights. 

According  to  the  work  of  Mangan(4)  and  of  Alezais(5), 
the  bird  has  an  average  kidney-to-body  weight  of  6.9 
grams  per  kilo  while  man  has  a  ratio  of  4.3  grams,  dog  5.9 
grams  and  guinea-pig  8.5  grams.  The  first  author  would 
show  that  the  fisheating  birds  have  the  heaviest  and  vege- 
tarian birds  the  lightest  organ.  Our  own  figures  are 
limited  to  the  weights  of  apparently  normal  organs  in 
thirty-one  mammals  and  five  birds ;  they  are  as  follows : 

per  kilo  of  body  weight 


Primates (5) 

7.7  grams 

Carnivora ....  (6) 

7.6  grams 

Rodentia (2) 

15.    grams 

Hyracoidea.  .  .(1) 

7.5  grams 

Ungulata (9) 

3.5  grams 

Edentata (1) 

5.6  grams 

Marsupialia..  .(7) 

7.6  grams 

Monotremata.(l) 

11.2  grams 

Average  7. 

Steganopodes .  (1) 

9.1  grams 

Anseres (1) 

3.9  grams 

Struthiones...(3) 

7.    grams 

Average  6.7 

This  to  be  sure  is  not  a  very  exhaustive  list  but  is  the 
result  of  our  routine  observations  and  subject  to  all  limi- 

(4)  Bull  Mtts.  Hist.  Xat.,  1911,  493  and  1912,  527,  and  C.  R.  Acad. 
8c.,  155,  182. 

(5)  C.  R.  Soc.  Biol,  1898,  5,  188. 

18 


266  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

tations  of  such  work.  Grossly  diseased  organs  are 
naturally  excluded.  More  avian  weights  are  not  available 
because  of  the  difficulty  of  removing  the  organ  from  its 
bed,  in  a  manner  assuring  us  of  completeness.  I  am 
inclined  to  view  our  mammalian  records  as  fairly  repre- 
sentative. Figures  to  be  found  in  text-books  of  human 
and  veterinary  anatomy  correspond  to  those  given  by  the 
authors  just  quoted  and  in  our  own  list.  The  values  for 
rodents,  hyraces,  edentates  and  monotremes  may  be  modi- 
fied by  more  figures. 

There  is  however  one  point  which  does  not  appear  in 
the  list.  Small  animals  have  relatively  larger  kidneys 
than  large  animals.  Tliis  is  perhaps  most  strikingly 
illustrated  among  the  ungulates  which  have  the  lowest 
value  quoted.  A  small  deer  had  a  kidney-to-body  index 
of  5.9  grams  per  kilo  while  a  camel  had  only  2.8.  Judging 
by  the  work  of  Magnan  the  avian  kidney  should  be  larger 
/than  the  mammalian,  a  conclusion  with  which  I  am 
inclined  to  coincide,  even  though  the  weights  cited  do  not 
bear  this  out. 

In  so  far  as  the  function  and  chemistry  of  the  kidney 
and  its  excretion  are  concerned  this  study  can  supply 
little.  The  general  metabolism  is  known  for  most 
animals,  it  being  dependent  upon  diet  and  gastrointestinal 
discharge  of  excrement.  What  lessons  can  be  learned 
will  be  discussed  by  Dr.  Corson- White  in  the  section  on 
diet.  Our  observations  upon  the  ability  of  the  kidney  to 
excrete  normal  urine  are  limited  to  the  examination  of 
vesical  contents  at  death  or  of  the  occasional  specimen 
obtained  in  cages  in  the  quarantine  room.  Renal  disease 
was  formerly  considered  of  little  or  no  importance  in 
veterinary  medicine  or  at  least  was  studied  only  as  a 
specific  separate  and  occasional  occurrence.  Kitt(6) 
systematized  the  knowledge  of  the  subject  at  the  time  he 
wrote  but  it  remained  for  Hutyra  and  Marek  in  their 
text-book  to  emphasize  its  general  importance  and  to 

(6)   Monatsh.,  1893. 


THE  URINARY  TRACT  267 

clarify  diagnostic  measures.  Breindl(7)  pointed  out  that 
nephritis  occurs  more  often  in  acute  general  diseases, 
notably  the  specific  infections,  than  was  customarily 
thought,  thus  placing  the  subject  for  the  lower  animal 
where  it  is  in  human  medicine. 

Renal  disease  is  quite  common  among  wild  animals 
albeit  there  are  certain  orders  in  which  the  lesions  are 
less  conspicuous.  Clinical  diagnoses  of  nephritis,  ajid 
this  is  the  only  diagnosis  attempted,  have  been  made  on 
monkeys  by  examination  of  urine  which  shows  the  same 
characters  as  in  the  human  disease.  In  ungulates  more 
attention  is  to  be  placed  upon  the  cellular  contents  of  the 
urine  since  renal  epithelium  is  apparently  shed  more 
readily  and  casts  less  often  formed.  Signs  and  symptoms 
of  renal  disease  are  limited  to  edema  and  uremia;  cases 
of  the  latter  are  rare  enough  to  discuss  separately  at  the 
proper  place. 

Absence  of  Abnormalities. 

Abnormalities  of  size,  shape  and  position  of  the  kidney 
are  frequently  reported  in  literature  of  veterinary  medi- 
cine and  aplasia  has  been  described.  Our  material  has 
failed  to  present  cases  of  horse-shoe  kidney  well  known  to 
occur  in  horses,  cows,  sheep  and  dogs.  Wandering 
kidneys  are  also  known  but  have  not  been  seen  in  our  wild 
animals.  Shall  these  abnormalties  be  considered  as  due 
to  degenerative  changes  in  cross  bred  animals  or  as  the 
result  of  the  strain  of  domestication?  To  such  a  specula- 
tive question  our  material  affords  no  answer. 

Hypertrophy. 

That  the  kidney  has  the  power  of  hypertrophy  in  a 
compensatory  manner  is  illustrated  by  two  cases.  A 
Japanese  Macaque  {Macacus  fuscatus)  6  apparently  had 
suffered  with  a  unilateral  nephritis  which  had  gone  into  a 
contracted  stage.  At  all  events  much  functionating  tissue 

(7)  Inaug.-Diss.  Oiessen,  1911. 


268   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

was  gone,  the  organ  irreg-ular  and  small,  being  half  or  less 
of  the  size  of  the  other  organ  which  was  larger  than  is 
considered  normal  for  the  species.  Histologically  the 
large  organ  was  practically  normal.  A  common  opossum 
{Didelphys  virginiana)  suffered  mth  a  complete  suppura- 
tive nephritis  of  the  right  side  which  completely  destroyed 
the  organ;  the  origin  of  this  is  not  clear  as  no  ascending 
disease  could  be  determined  and  no  certain  acute  infection 
had  existed;  decomposition  precluded  satisfactory  bac- 
teriology. The  left  kidney  was  nearly  twice  its  normal 
size  and  involved  in  an  early  diffuse  nephritis,  with 
miliary  abscesses,  in  which  the  glomeruli  did  not  partici- 
pate. There  were  in  these  sections  evidences  of  regenera- 
tion, swollen  reduplicated  epithelial  coverings  presenting 
a  picture  similar  to  those  seen  in  so-called  chronic 
nephritis  secondary  to  interstitial  change. 

Infiltrations. 

Pathological  infiltrations  of  the  renal  structures  are 
exceedingly  uncommon.  Early  in  our  experience  we  were 
often  perplexed  at  the  appearance  of  certain  organs, 
notably  in  carnivores  and  marsupials  to  which  we  were 
inclined  to  apply  the  term  fat  infiltration.  However  the 
absence  of  reasons  for  considering  this  picture  pathologi- 
cal seemed  sufficient  cause  to  ignore  the  finding,  and  later 
Pfeiffer(8)  called  attention  to  the  apparent  inability  of 
these  kidneys  to  emulsify  fat  or  at  least  to  combine  it  in 
an  invisible  form,  an  ability  possessed  by  the  herbivorous 
organ.  A  monkey  and  a  passerine  bird  only  showed  suffi- 
cient fat  visible  in  the  renal  epithelium  to  warrant  a 
denomination  of  fatty  infiltration ;  these  were  both  obese 
specimens.  Amyloid  infiltration  occurred  in  four  mam- 
mals and  six  birds,  being  a  sequel  of  its  usual  causes, 
tuberculosis,  chronic  suppuration  and  osseous  system 
disease.  It  is  perhaps  well  to  emphasize  the  fact  that 
every  organ  the  seat  of  amyloid  deposit  need  not  be 

(8)   Arch.  f.  Tierheilk.,  V.  38-99. 


THE  URINARY  TRACT  269 

enlarged.  This  teaching  is  common  but  I  have  seen  human 
cases  without  enlargement  and  only  two  of  the  ten  cases 
in  these  animals  are  noted  as  bulkier  than  normal. 

Degenerations. 

Degenerations  represent  the  reaction  of  the  Iddney  to 
toxic  or  infectious  agents  and  might  be  considered  as  indi- 
cating the  vulnerability  of  the  organ.  Their  incidence 
does  not  coincide  with  that  of  nephritis  as  we  shall  see 
later.  Any  discussion  of  degenerative  phenomena,  and 
especially  in  the  kidney,  should  be  limited  by  a  definition 
of  what  they  are  believed  to  be  and  their  separation  from 
inflammations.  Degenerations  are  swellings,  granular- 
ities, vacuolizations  or  infiltrations  of  tubular  epithelium, 
changes  which  destroy  the  outline  and  internal  structure, 
perhaps  including  the  nucleus.  No  changes  of  the 
glomerular  tuft  or  interstitial  tissue  are  necessary  for 
this  conception  since  when  these  occur  the  picture 
becomes  that  of  nephritis.^  In  border-line  cases  it  is 
safer  to  include  the  case  under  the  latter  heading  since 
then  the  physiology  is  apt  to  be  disturbed,  albumen  and 
casts  appearing  in  the  urine.  Degenerations  appear  in 
various  pathological  states — toxemia,  infection,  prolonged 
congestion  and  others.  The  first  named  cause  seems  to  be 
the  most  important  in  our  records  and  the  seat  of  the 
toxine  production  seems  to  be  the  intestine.  Enteritis 
stands  very  high  in  the  list  of  accompanying  factors,  espe- 
cially in  Carnivora,  Primates  and  in  Aves.  Perhaps  the 
most  instructive  cases  are  to  be  found  in  the  Ungulata 
with  toxic  duodenitis.  The  kidney  in  these  animals  is 
deep  red  or  purple,  with  a  spanned  capsule.  The  section 
surface  bulges  slightly,  is  of  an  opaque,  dull  purple  color 
and  shows  a  congested  zone  between  cortex  and  medulla. 
Tubular  epithelium  may  be  found,  under  the  microscope, 
sufficiently  swollen  to  fill  the  lumina,  in  places  being  like 
ground  glass,  in  others  distinctly  vacuolated.    It  cannot 


270   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

be  stated  absolutely  which  part  of  the  tubule  is  usually 
affected;  it  seems  more  often  the  distal  convoluted  por- 
tions than  other  subdivisions.  Glomeruli  may  be  full  of 
blood  but  there  is  no  increase  in  cell^  nor  any  material  in 
the  capsular  space.  The  urine  is  dark  and  may  or  may 
not  show  albumen.  The  kidney  of  kangaroos  with  strepto- 
thricosis  is  similar  to  the  picture  just  given. 

Mammals  have  sho^vn  a  percentage  incidence  of  renal 
degenerations  of  4.8  per  cent,  wliile  Aves  show  only  3,  per 
cent.  In  order  of  incidence  the  carnivores  head  the  list 
followed  by  Lemures,  Accipitres,  Rodentia,  Primates  and 
Marsupialia;  the  remaining  groups  show  but  a  few  cases. 
(  A  form  of  degeneration  is  sometimes  seen  in  the  a\ian 
kidney  the  seat  of  excess  urate  collections,  especially 
when  these  are  arranged  as  so-called  uratic  infarcts.  This 
last  term  has  been  applied  to  the  streaking  and  mottling 
of  human  kidneys  by  the  accumulations  of  these  salts  in 
a  manner  believed  by  some  to  be  related  to  the  formation 
V  of  uratic  calculi.  The  epithelium  of  such  a  kidney  may 
show  granularity  and  collections  of  acids  and  salts  have 
been  found  in  the  lumina.  In  the  bird  on  the  other  hand 
one  frequently  sees  masses  of  urates  in  one  lobule,  or  a 
part  thereof,  arranged  to  simulate  closely  the  common 
infarct  shape.  Secretory  cells  in  the  affected  area  are 
hydropic,  with  absent  or  dislocated  nucleus,  or  again  they 
present  a  densely  basic-staining  protoplasm.  This  form 
of  kidney  is  well  seen  in  what  has  been  called  here  an 
uratic  serositis,  a  coating  of  all  somatic  free  membranes 
■\\^th  a  thin,  white,  granular  film.  We  have  tried  with 
many  teclmiques  to  preserve  one  of  these  cases  but  the 
deposit  either  dissolves  or  the  whole  specimen  becomes 
opaque.  Although  the  term  infarct  is  applied  to  these 
lesions,  they  are  of  course  not  infarcts  in  the  customary 
use  of  the  term.  True  infarcts  are  exceedingly  uncommon  j 
and,  with  the  exception  of  one  case  which  became  infected 
and  suppurated,  have  been  negligible  in  our  material. 


THE  URINARY  TRACT  271 

Hemoerhages. 

Hemorrhages  into  the  kidney  are  found  in  acute  infec- 
tions and  certain  diseases  like  leucemia;  they  are  of  little 
moment.  Perirenal  hemorrhage  is  a  somewhat  striking 
and  unusual  affair.  Recently  I  saw  at  a  human  autopsy 
of  a  young  subject  a  subcapsular  hemorrhage  from  the 
renal  substance  probably  due  to  vascular  rupture  in  an 
acute  nephritis;  there  was  no  history  of  injury.  There 
have  been  three  cases  of  subcapsular  hemorrhage  in  our 
records  and  as  two  of  them  represented  the  immediate 
cause  of  death,  are  interesting  enough  to  record.  An 
armadillo  suffered  an  acute  diffuse  nephritis  mth  much 
congestion  but  not  enough  to  call  it  hemorrhagic.  There 
was  a  large  hemorrhage  around  the  left  organ,  probably 
from  a  vessel  near  the  hilum,  sufficient  to  compress  the 
kidney  and  cause  it  to  atrophy.  A  lion  presented  an 
acute  vegetative  endocarditis  with  all  its  usual  compli- 
cations. The  right  renal  capsule  was  distended  with 
recent  clot  to  a  size  which  reached  to  the  pelvic  brim. 
Presumably  an  embolism  caused  thrombosis,  ulceration 
and  rupture  of  some  middle  size  vessel.  A  dormouse  suf- 
fering with  an  acute  general  infection  probably  emanat- 
ing from  the  intestine,  had  several  small  recent  clots 
separating  the  kidney  from  its  capsule. 

Nephritis. 

Nephritis,  whether  one  begin  its  conception  with  the 
clinicopathological  picture  originally  given  by  Bright, 
with  the  purely  pathological  classification  of  Weigert  and 
Virchow  or  the  modern  tendency  to  subordinate  all  physi- 
cal changes  to  clinical  phenomena,  is  nevertheless  a 
process  of  degeneration  and  inflammation  affecting  the 
secreting  and  supporting  structures  of  the  kidney  and 
leading  to  some  degree  of  impaired  function.  The  disease 
is  bilateral  in  so  nearly  every  case  that  for  practical 
purposes  unilateral  cases  may  be  ignored.  This  implies 
that  for  some  reason  the  renal  tissues  are  generally  sus- 


272   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

ceptible  to  etiological  agents  so  that  when  one  side  is 
affected  its  fellow  seems  always  to  participate  or  to 
follow.  It  seems  desirable  in  studying  nephritis  to  eval- 
uate fully  the  mutual  relations  of  functionating  and  sup- 
porting tissues  and  of  the  various  sections  of  the  first 
named.  It  is  taught  in  many  places  that  inflammations 
of  one  or  another  of  these  parts  may  occur  independently, 
as  for  example  a  tubular  nephritis,  a  glomerulonephritis 
and  an  interstitial  nephritis.  If  however  one  reflect  upon 
the  dependence  of  the  tubular  function  upon  the  glomer- 
ulus and  vice  versa  or  upon  the  effect  of  inflammatory 
exudates  in  the  supporting  tissues  upon  the  blood  supply 
of  the  tubule,  it  becomes  evident  that  only  the  most 
trivial  or  evanescent  pathological  changes  in  one  can  be 
without  effect  upon  the  others.  It  is  diflBcult  to  see  how, 
for  examples,  a  glomerulus  could  remain  normal  if  its 
associated  tubule  were  destroyed  or  how  if  round  cell 
infiltration  or  pus  surround  a  capsule  for  any  length  of 
time,  this  structure  could  fail  to  be  doomed.  All  this  is 
by  way  of  directing  attention  to  the  progress  of  physi- 
cal damage  in  a  kidney  which  has  received  injury  sufficient 
to  cause  nephritis,  but  of  course  it  does  not  explain 
the  cause. 

In  classification  of  nephritis  different  commentators 
have  employed  different  standards  according  as  they 
viewed  the  acuteness  or  chronicity  of  the  process,  or  as 
the  principal  functionating  structures,  glomerulus,  tubu- 
lar epithelium,  blood  vessels,  or  supporting  connective 
tissues,  presented  the  most  conspicuous  changes.  To 
these,  clinicians  have  added  phenomena  of  constitutional 
complication  or  of  direct  renal  insufficiency.  These  latter 
being  unavailable  for  us,  we  must  fall  back  upon  a 
classification  based  upon  physical  changes  and  to  this  end 
we  have  always  used  a,  slight  modification  of  the  Weigert 
method.  This  classification  offers  little  in  the  direction 
of  etiology  except  that  toxins  are  believed  to  cause 
tubular  changes,  bacteria  to  produce  glomerular  lesions 


THE  URINARY  TRACT  273 

and  vascular  deficiencies  to  lie  at  the  root  of  chronic 
interstitial  nephritis. 

The  origin  of  acute  nephritis  of  chiefly  degenerative 
character  seems  best  explained  by  reference  to  some  form 
of  toxemia,  whereas  exudative  processes,  be  they  in 
glomerulus  or  supporting  structures,  seem  to  depend 
upon  the  direct  action  of  bacteria.  The  origin  of  a  chronic 
nephritis  cannot  be  explained  quite  so  readily.  No  one 
has  answered  with  complete  satisfaction  whether  a 
chronic  process  always  begins  with  and  proceeds  from  a 
single  attack  of  acute  disease,  whether  many  acute 
attacks  succeed  upon  one  another  or  whether  many  small 
crops  of  agents  successively  attack  the  organ  over  a  long 
time.  Nor  has  an  adequate  explanation  of  the  role  of 
damaged  blood  vessels  been  given.  It  is  reasonably  easy 
in  man  to  discover  the  existence  of  nephritis  and  of  a 
possible  cause;  this  is  only  true  of  acute  cases  in  wild 
animals.  Focal  infections,  those  which  might  be  the  point 
of  mobilization  for  bacteria  sent  to  the  kidneys,  are  fre- 
quently found  in  man  but  mth  exception  of  an  occasional 
carious  tooth,  or  a  chronic  osteitis  are  to  be  localized  with 
difl&culty  in  lower  animals.  In  so  far  as  the  role  of  a  single 
acute  attack  in  the  causation  of  chronic  disease  is  con- 
cerned our  material  offers  nothing,  but  some  collateral  or 
presumptive  evidence  may  be  mustered  in  regard  to 
multiple  infections. 

Wild  animals  do  not  give  evidence  of  repeated  attacks 
of  acute  disease  and  indeed  it  would  seem  that  they  more 
often  die  of  an  acute  infection  than  live  to  have  it 
repeated.  Evidences  of  chronic  infection,  not  focal,  are 
reasonably  definite  in  fortj^-eight  per  cent,  of  the  cases 
of  chronic  nephritis  encountered  here.  This  suggests 
strongly  that  in  this  material  protracted  infectious  states 
offer  opportunities  for  renal  damage  of  progressive  char- 
acter. Vascular  disease  has  been  found  twenty-eight 
times  (see  also  section  on  arteries),  in  twenty-six  of  which 
the  nephritis  seemed  due  to  or  advanced  by  the  damage  to 


274  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

the  vessels.  This  means  further  that  only  14,3  per  cent,  of 
the  chronic  forms  and  4  per  cent,  of  the  total  seem  closely 
related  to  disease  of  blood  vessels. 

Nephritis  has  been  found  in  12.2  per  cent,  of  our  total 
autopsies.  Mammals  show  an  incidence  of  20.6  per  cent., 
birds  7.7  per  cent.  Only  the  orders  upon  which  more  than 
one  hundred  autopsies  have  been  held  are  subjected  to 
separate  analysis.  Some  of  the  remaining  orders  give 
very  high  figures  which  may  indicate  great  renal  \nilner- 
ability  but  it  is  deemed  unfair  to  make  statements  upon 
them/  Carnivorous  mammals  and  birds  lead  their  respec- 
tive classes,  the  succeeding  order  of  renal  vulnerability 
being  marsupials,  ungulates,  rodents.  Primates,  .Galli, 
Striges,  anserine  birds,  parrots,  and  doves.'  The  leaders 
of  this  list,  Carnivora  and  Accipitres,  occupy  a  definite 
position  in  the  analysis  of  acute  and  chronic  lesions. 
Their  kidneys  show  the  smallest  percentages  of  acute 
lesions  and  the  highest  percentages  of  chronic  lesions. 
This  would  seem  to  indicate  a  resistance  to  acute  injuries 
but  susceptibility  to  prolongated  or  repeated  infections  or 
intoxications.  The  relation  of  chronic  infection  of  some 
sort  to  chronic  renal  disease  is  not  as  clear  as  the  influence 
of  acute  infection  to  acute  nephritis.  Taking  Carnivora 
for  example  with  34.2  per  cent,  of  chronic  nephritis  we 
find  22.4  per  cent,  with  evident  chronic  inflammation  while 
in  the  55  per  cent,  of  acute  forms  40  per  cent,  are  of  acute 
infectious  origin — the  relation  is  as  64  is  to  74. 

While  the  relation  of  infection  to  nephritis  is  a  con- 
sistent and  perfectly  acceptable  one,  the  frequency  of  this 
disease  in  the  carnivorous  orders  obliges  one  to  think  of 
high  protein  diet  as  a  favoring  factor.  Renal  disease  is 
conmion  enough  in  other  orders,  some  strictly  herbiv- 
orous, and  it  is  fair  only  to  emphasize  which  are  the 
leaders  in  incidence.  In  so  far  as  anatomy  or  habits  are 
concerned  no  generalizations  seem  permissible.  There  is 
no  relation  of  nephritis  to  the  size  of  the  kidney  as  given 


THE  URINARY  TRACT  275 

on  a  previous  page,  to  the  length  of  the  alimentary  tract, 
or  to  the  expected  longevity. 

Toxic  nephritis  is  a  term  applied  when  the  kidney  is 
the  seat  of  epithelial  degeneration,  much  congestion,  per- 
haps leading  to  tiny  hemorrhages,  and  definite  swelling  of 
the  tuft  without  exudation  into  the  capsular  space.  It  is 
a  severe  grade  of  the  degenerations  already  mentioned 
and  is  exemplified  by  the  organ  in  cases  of  acute  duode- 
nitis of  ungulates  and  in  some  monkeys  dying  after 
tuberculin  injection.  It  seems  especially  to  follow  gastro-  ^ 
intestinal  diseases  believed  to  be  due  to  food  intoxications. 
It  seems  important  in  monkeys  and  wild  rodents.  No 
adequate  explanation  is  at  hand  for  the  latter. 

As  has  already  been  stated  vascular  disease  was 
present  in  twenty-six  cases  in  a  manner  suggesting  some 
relation  to  the  cardiorenal  complex  but  the  only  organ  to 
which  the  term  renal  sclerosis  of  arteriosclerotic  origin 
could  be  applied  is  that  of  an  eagle ;  the  autopsy  is  cited. 

Bald  Eagle  {Haliceetus  leucocephalus) .  $  General  obliterating  en- 
doarteritis.  Chronic  interstitial  nephritis.  Passive  congestion  of  liver. 
Chronic  localized  myocarditis.  Near  the  apex  of  the  heart  the  muscle 
shows  a  slight  opacity.  The  kidney  is  enlarged,  firm,  section  surface 
glistening.  Both  section  and  surface  show  a  mottled  brown  and  white 
appearance,  following  particularly  on  section  the  division  into  cortices 
and  medidlge.  Digestive  system  apparently  normal.  Microscopic  sec- 
tion of  heart  muscle  from  the  wall  of  the  ventricle  shows  well  preserved 
muscle  fibres  with  a  slightly  unusual  degree  of  pigmentation.  Section 
from  valve  base  shows  a  definite  interfascicular  and  intraf aseicular  fibro- 
sis which  is  co-extensive  with  a  similar  thickening  of  the  endo-  and 
pericardium.  The  new  tissue  under  the  latter  is  edematous.  The  valve 
itself  is  thickened  the  fibres  swollen  and  hyaline.  There  is  no  reduplica- 
tion of  the  endothelium.  One  artery  in  the  muscle  is  obliterated.  This 
is  not  associated  with  any  degeneration  of  the  muscle  in  the  section. 
Section  of  kidney  shows  the  pale  areas  noted  grossly  to  be  made  up 
of  groups  of  arteries  with  their  extensive  coalescing  adventitise.  The 
changes  in  the  arteries  are  precisely  the  same  as  those  seen  in  the  liver 
but  are  more  extensive.  Connective  tissue  goes  out  from  the  arteries 
into  the  parenchyma  distorting  the  tubules  and  enclosing  the  glomeruli 
so  that  the  capsule  of  the  latter  is  much  thickened.  Epithelium  is 
granular,  in  some  places  absent,  nearly  always  low.  Section  of  liver 
shows  general  parenchyma  practically  normal  with  slight  granularity 
in  places  and  moderate  passive  congestion.     Veins  are  negative  but 


276   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

arteries  show  a  general  arteritis.  The  picture  varies  somewhat  in  differ- 
ent arteries  ranging  from  a  simple  thickening  of  the  adventitia  to  a 
change  involving  all  three  layers.  There  is  hyaline  change  in  the  media  in 
many  sections.  Lumen  is  in  all  cases  reduced  and  in  some  there  is 
active  intimal  proliferation  in  excess  of  what  would  be  expected  in 
connection  with  the  medial  change.  A  few  of  the  arteries  have  their 
lumen  completely  obliterated. 

This  is  meant  to  illustrate  the  picture  of  vascular 
disease  in  the  kidney  in  the  absence  of  satisfactory  evi- 
dence that  nephritis  per  se  antedated  or  accompanied 
changes  in  the  vessels.  In  such  cases  vascular  disease 
dominates,  renal  parenchymatous  damage  being  rela- 
tively inconspicuous.  Two  old  carnivores,  a  paradoxure 
and  a  skunk,  presented  shrunken  kidneys  \vith  prominent 
wide-walled  vessels  but  in  these  some  definite  evidence  of 
old  nephritis  was  at  hand. 

In  so  far  as  the  relation  of  senility  to  nephritis  is  con- 
cerned the  data  at  hand  are  not  conclusive.  In  many  old 
animals  some  degree  of  fibrosis  is  present  mthout  the 
existence  of  truly  destructive  changes  in  the  parenchyma. 
Plimmer  of  London  writes  that  there  is  increased  nephri- 
tis in  old  age  but  from  our  material  I  would  be  inclined 
to  put  in  that  in  many  cases  the  nephritis  was  the  reason 
for  old  age  rather  than  that  old  age  brought  on  a 
nephritis.  However  the  exact  length  of  life  and  of 
captivity  is  known  in  too  few  specimens  to  make  a  con- 
clusion justified. 

Ascending  Nephritis. 

There  is  some  difference  of  opinion  as  to  the  definition 
of  the  term  ascending  nephritis,  a  confusion  arising  partly 
from  the  intended  meaning  of  the  participial  adjective, 
partly  from  the  frequency  with  which  infections  or 
obstructions  of  the  urinary  outlets  antedate  or  accompany 
suppurative  nephritis.  Perhaps  our  records  may  help  to 
straighten  out  this  matter. 

Ascending  nephritis  means  for  our  study  an  infection 
which  passes  from  the  pelvic  surfaces  of  the  pyrainids 


THE  URINARY  TRACT  277 

outward  toward  the  renal  capsule.  Thus  it  is  immaterial 
whether  there  be  or  be  not  an  obstruction  lower  down. 
Such  forms  of  nephritis  are  infiltrative,  frequently  puru- 
lent and  are  dependent  upon  pyelitis  or  the  settling  of 
bacteria  in  the  deepest  parts  of  the  medulla. 

Three  explanations  are  given  in  human  medicine  for 
the  origin  of  this  lesion.  Some  observers  assume  a  direct 
transmission  of  bacteria  up  the  ureter  from  an  infected 
bladder  or  urethra,  in  a  direction  contrary  to  the  urinary 
current.  This,  it  is  believed  by  some  can  occur  only  in  the 
presence  of  physical  obstruction,  stone,  kink,  or  pressure 
of  adjacent  masses  upon  the  ureter,  whereby  its  blood 
supply  is  damaged  and  infection  facilitated.  Others 
would  explain  the  path  of  infection  as  the  lymphatics  of 
the  ureteral  wall  which  are  infected  at  the  opening  in  the 
bladder  by  a  deep  seated  cystitis  or  by  infection  from  a 
periproctitis  or  from  the  female  genitalia.  Still  another 
explanation  is  offered  by  those  who  do  not  credit  ascend- 
ing infections.  They  would  have  it  that  pyelitis  arises 
from  bacteria  in  the  blood  stream  and  only  in  the  presence 
of  injury  (calculus),  ureteral  dilation  (slowly  progressive 
stenosis,  or  kinks,  floating  kidney,  pressure  by  pregnant 
uterus)  and  similarly  operative  factors.  There  should 
be  excluded  from  tliis  category  cases  of  chronic  nephroli- 
thiasis, and  of  stone  only  in  the  pelvis.  Under  such  con- 
ditions it  is  inevitable  that  a  low  grade  of  fibrosis  with 
damage  to  the  secretory  structures  should  exist,  even  in 
the  absence  of  active  bacterial  invasion.  The  cases  are 
only  important  for  our  present  subject  when  active 
bacterial  infection  is  implanted  upon  them. 

Among  our  autopsies  there  have  been  found  fourteen 
cases  of  ascending  nephritis  and  seven  cases  of  pyelitis ; 
the  most  instructive  examples  are  mentioned  briefly  as 
follows :  !  Five  of  this  twenty-one  were  associated  with 
general  infectious  diseases  (three  septicemias)  and  pre- 
sented no  evidence  of  ascending  obstruction.  Two  of  this 
five  were  a  bear  and  a  fox,  the  former  suffering  with 


278  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

streptothricosis  septicemia,  the  latter  with  distemper; 
one  was  a  fox  whose  pelvic  and  renal  lesion  seemed 
entirely  primary  for  no  apparent  focus  was  detected ;  two 
were  birds  with  acute  general  infection.  These  cases  seem 
therefore  to  be  instances  of  primary  pyelitis.  The  follow- 
ing group  includes  cases  with  inflammation  low  in  the 
urinary  tract.  A  fox  had  a  cystitis,  urethritis  and  colitis, 
a  distinct  edema  and  congestion  being  found  in  the  pelvis 
around  the  rectum.  Two  opossums  had  cystitis,  one  due 
to  a  traumatic  urethritis,  the  other  secondaiy  to  a  pros- 
tatitis of  undetermined  causation.  A  raccoon  had  a 
chronic  cystitis  with  swelling  and  edema  of  the  first  parts 
of  ureter.  A  parrakeet  showed  ureteritis,  pyelitis  and 
nephritis  from  simple  cloacitis. 

Where  obstruction  was  more  definite  the  following 
cases  were  observed.  Suppurative  nephritis  succeeded 
upon  gangrenous  cystitis  after  uterine  prolapse  in 
a  deer.  The  following  cases  of  pyelitis  and  nephri- 
tis were  associated  with  calculus,  only  in  the 
renal  pelvis4-an  armadillo,  Tasmanian  devil,  a 
deer  and '  a  goose.  A  cockatoo  had  a  stone  in  the 
cloaca  which  seemed  to  cause  a  definite  obstruction  to 
both  ureters  and  a  catarrhal  inflammation  of  the  wall. 
Pyelitis  and  interstitial  nephritis  can  also  follow  exces- 
sive urate  collections  in  birds.  At  a  later  time  this  will  be 
discussed  more  fully,  but  at  this  place  two  cases  of  dis- 
tinct abscess  formation  in  a  renal  lobule  based  upon 
urate  collections  may  be  mentioned  since  in  a  measure 
the  lesions  were  dependent  upon  obstruction. 

It  is  therefore  evident  that  all  the  theories  of  the 
causation  of  pyelitis  with  resultant  nephritis  seem  accept- 
able. It  has  been  claimed  that  bacteria  may  be  found  in 
the  blood  stream  before  evidences  of  pyelitis  present 
themselves.  Concerning  this  our  records  offer  no  infor- 
mation, but  it  is  worthy  of  note  that  five  of  twenty-one 
instances  gave  a  picture  of  septicemia.  It  is,  however, 
fair  to  state  that,  while  mild  cases  of  pyelitis  occurred 


THE  URINARY  TRACT  279 

where  there  was,  judging  from  the  protocol  and  histo- 
logical sections,  obvious  opportunity  for  its  ascent,  in  ten 
other  cases  of  cystitis  and  urethritis  no  pelvic  or  renal 
disease  is  recorded;  two  of  these  were  acute  exudative 
cystitis  and  one  was  a  tumor.  It  seems  that  wild  animals 
seldom  live  long  enough  to  have  obstructions  exert  back 
pressure  of  urine  to  the  extent  which  one  is  accustomed 
to  see  in  human  medicine.  Hydroureter  and  hydro- 
nephrosis have  not  been  seen. 

Abscess  of  the  kidney  has  occurred  occasionally  in  the 
metastatic  form  and  only  twice  as  the  large  destructive 
process  such  as  is  seen  in  the  human  being  (surgical 
kidney).  One  massive  abscess  was  seen  involving  about 
one-third  of  the  organ,  and  this  seems  to  have  had  a 
tuberculous  basis.  Another  destructive  purulent  nephri- 
tis was  quoted  on  page  268  when  illustrating 
compensatory  hypertrophy. 

Examination  of  records  and  preserved  specimens  of 
nephritis  reveals  few  striking  differences  which  might  be 
considered  characteristic  for  the  various  orders.  This 
is  possibly  due  to  the  fact  that  the  lesions  have  been 
classified  under  the  same  system,  a  method  which  has 
proven  convenient  and  consistent.  In  support  of  this 
one  might  refer  to  that  form  of  nephritis  which  gives  the 
most  definite  clinical  and  pathological  picture  of  renal 
disease,  namely  the  chronic  parenchymatous.  This  is 
fairly  weU  represented  in  Primates,  Carnivora  and  Ungu- 
lata.  In  one-fourth  of  the  cases  one  finds  distinct  edema, 
especially  in  the  body  cavities,  cervical  and  mediastinal 
tissues,  and  in  one-fifth  an  appreciable  grade  of  anemia; 
uremia  was  the  terminal  picture  in  one  animal  of 
each  order. 

Histology  of  Nephritis. 

An  attempt  to  discover  minute  lesions  peculiar  to  the 
various  groups  gives  results  that  are  far  from  satisfac- 
tory.    With  reserve  it  may  be   said  that  carnivorous 


280   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

animals  show  a  tendency  to  greater  interstitial  and 
glomerular  lesions  than  do  herbivorous  ones,  and  that 
casts  are  more  often  found,  in  all  kinds  of  nephritis,  in 
the  former  varieties.  Rodents  are  conspicuous  excep- 
tions to  this  statement,  since  they  frequently  have 
glomerular  lesions  and  casts  in  abundance ;  this  exception 
exemplifies  the  unwisdom  of  drawing  definite  conclusions 
in  this  respect. 

Birds  as  contrasted  with  mammals  show  very  promi- 
nent tubular  and  inconspicuous  glomerular  lesions.  The 
principal  alterations  seen  in  the  avian  kidney  are  round 
cell  infiltrations  of  the  deep  cortical  and  outer  medullary 
zones,  and  cloudy  or  hydropic  swelling  of  the  convoluted 
tubules.  In  chronic  cases  intertubular  fibrosis  is  clear 
but  not  so  definite  as  the  perivascular,  while  the  glomer- 
ular tufts  are  occasionally  wholly  normal.  Wlien  these 
are  destroyed  it  seems  to  have  occurred  by  compression 
rather  than  by  inflammation.  A  form  of  nephritis  peculiar 
to  birds  might  be  termed  local  necrotizing.  It  seems  to 
be  due  to  local  urate  deposits  and  to  have  its  origin  like 
that  form  already  mentioned  under  acute  interstitial 
nephritis.  It  has  been  seen  in  avian  gout,  a  condition  in 
which  the  kidneys  may  or  may  not  have  visible  masses  of 
hardened  urates  in  them.  The  gross  picture  is  of  a  spotty 
pale  organ  of  a  gray-brown  color.  Minutely  studied  the 
medulla,  adjacent  cortical  tubules  and  perhaps  the  pelvic 
tissues  will  present  an  opaque  condition  taking  a  diffuse 
basic  dye.  Crystals  have  not  been  seen.  This  form  is 
especially  common  in  Columbae,  Psittaci  and  Herodiones. 

Geneeal  Effects  of  Nephritis. 

The  results  of  nephritis  generally  speaking  are  edema, 
anemia,  cardiac  hypertrophy,  inflammations  of  the  se- 
rous surfaces  and  uremia.  Aside  from  the  cases  of  chronic 
parenchymatous  nephritis  cited  above,  edema  has  been 
decidedly  inconspicuous.  It  may  be  found  in  avian  cases 
of  acute  and  subacute  nephritis,  about  the  flanks  and  in 


THE  URINARY  TRACT  281 

the  thoracic  areolar  tissues,  but  is  only  exceptionally  per- 
ceptible before  the  body  is  opened.  Anemia  is  almost 
never  extreme.  The  bone  marrow  although  mentioned  in 
but  few  histories,  seems  unchanged.  Cardiac  hypertrophy 
was  observed  ten  times  in  652  cases  of  nephritis,  twice  in 
460  acute  and  subacute  cases,  eight  in  192  chronic  cases. 
In  the  former  no  myocarditis  was  found,  in  six  of  the 
latter  it  was  found.  Clinical  and  pathological  experience 
teaches  that  serous  surfaces  are  frequently  inflamed 
during  a  nephritis.  Observations  on  our  material  coincide 
with  this  statement,  but  do  not  offer  an  explanation  of  it. 
The  figures  should  be  examined  for  mammals  and  for 
birds  separately,  since  the  serous  cavities  of  the  former 
are  closed,  separated,  and  protected,  whereas  a  close 
apposition  exists  between  the  serous  membranes  and  the 
lungs  in  birds,  an  arrangement  facilitating  infection  from 
without.  Nevertheless  the  bird  has  definitely  less  sero- 
sitis  accompanying  nephritis  than  does  the  mammal — 
4.4  per  cent.,  versus  8.6  per  cent.  In  the  former  class 
70  per  cent,  of  these  accompany  acute  nephritis  associ- 
ated with  acute  general  disease  while  only  45  per  cent,  of 
the  8.6  per  cent,  of  mammals  had  serositis,  acute 
nephritis  and  general  infection.  This  indicates  clearly 
that  mammalian  renal  disease  has  some  effect  upon  serous 
membranes  other  than  the  simple  participating  coinci- 
dence of  the  two  types  of  changes  during  an  acute  general 
infectious  disease.  No  one  kind  of  nephritis  was  espe- 
cially characterized  by  this  complication. 

Uremia,  except  under  the  best  clinical  conditions,  is  a 
term  to  use  with  caution.  I  have  seen  several  monkeys, 
a  few  marsupials  and  carnivores  and  an  occasional  bird 
in  a  dazed  ataxic  condition,  sometimes  exliibiting  an 
atypical  clonic  or  tonic  convulsion,  with  fixed,  rather 
small  pupils.  To  this  picture  I  have  applied  the  name 
uremia,  and  upon  several  occasions  have  found  a  severe 
grade  of  nephritis.  I  must  admit  having  failed  to  find 
renal  change,  however,  with  this  clinical  picture,  espe- 

19 


282   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

cially  in  the  carnivores  and  parrots,  animals  which 
''throw  fits"  at  times  without  apparent  good  reason.  In 
so  far  as  the  monkey  is  concerned,  I  am  satisfied  that  the 
picture  is  similar  to  that  seen  in  the  human  being.  The 
London  Garden  reports  in  1917  the  occurrence  of  uremia 
in  a  Mandrill  {Papio  maimon). 

Calculi. 

Calculi  are  well  known  in  the  renal  system  of  the 
domesticated  animals  so  that  it  is  not  surprising  to  find 
them  well  represented  all  through  the  lower  orders. 
Their  structure,  composition  and  effects  do  not  differ, 
however,  and  it  remains  only  to  point  out  their  distribu- 
tion. One  of  the  frequent  results  of  renal  and  pelvic 
lithiasis  is,  however,  missing  in  our  records,  namely 
hydronephrosis,  and  indeed  a  pressure  dilatation  of  the 
ureter  and  pelvis  has  not  been  observed  in  all  our  experi- 
ence. This  would  seem  to  be  explained  first  upon  the 
infrequency  of  stone,  of  obstructive  new  growths  and 
inflammatory  strictures  and  second  by  the  fact  that  lower 
animals  do  not  long  survive  conditions  which  would 
occasion  back  pressure  upon  the  kidneys. 

Definite  renal  and  pelvic  calculi  have  occurred  in  five 
Ungulata,  one  each  in  Edentata  and  Steganopodes.  To 
these  might  be  added  cases  of  uratic  sand  in  one  Car- 
nivora,  one  Marsupialia,  and  uratic  stones  of  large  size 
in  the  cloaca  of  two  Passeres  and  one  Accipitres.  Uratic 
collections  within  the  avian  kidney  have  been  mentioned 
and  will  be  discussed  later. 

According  to  written  descriptions  and  three  pre- 
served examples  the  calculi  in  four  of  the  ungulates  are 
mixed  urates  and  carbonates,  although  one  in  a  deer  was 
said  to  be  "mulberry"  in  surface.  The  specimens  at  hand 
are  moderately  hard  with  rough  irregularly  crystalhzed 
outer  shells  like  carbonate  deposits.  Incomplete  moulding 
to  the  calyx  is  found.  The  pelvic  cavity  while  seeming  to 
be  enlarged  is  not  distended  nor  do  the  stones  assume  the 


THE  URINARY  TRACT  283 

*' antler"  character  and  distort  the  pyramid.  All  the 
animals  showing  these  stones  have  but  a  single  pyramid 
in  the  kidney.  The  fifth  example  in  the  ungulate  had 
early  stones  forming  in  the  apex  of  the  pyramid  and  not 
yet  discharged  into  the  pelvis.  Four  of  these  cases 
showed  stones  on  both  sides,  one  only  in  the  left  kidney. 

The  case  among  the  Edentata  concerned  an  Armadillo 
{Tatu  novemcinctus),  in  the  kidney  of  which  there  were 
definite  smooth,  round,  hard  pebbles  in  the  right  pyramid 
and  several  fine  grains  in  the  pelvis.  There  was  a  recent 
hemorrhage  in  the  pelvis  which,  from  local  appearances 
and  autopsy  survey,  was  due  to  the  lithiasis. 

A  Gannet  (Sula  hassana)  represents  the  only  avian 
true  calculus.  In  this  case  many  small,  hard,  yellow 
stones  occupied  the  right  pelves,  which  contained  also 
loose  urates.  The  lobules  were  much  distorted  and 
showed  a  mild  interstitial  change.  This  accumulation 
was  confined  to  the  left  side.  These  well  developed  cases 
having  been  mentioned  the  next  most  important  may 
be  quoted : 

Tasmanian  Devil  {Sarcophilus  ur sinus).  Inactive  and  rather  on  the 
decline  for  several  months.  Chronic  diffuse  nephritis  with  acute  inter- 
stitial exacerbation.  Calculi  in  kidney  pelvis.  The  kidney  size  is 
normal,  shape  irregular,  capsule  smooth,  strips  easily  leaving  a  rough 
mottled  green-brown  surface.  Consistency  is  soft,  tough,  resilient.  Cor- 
tex slightly  wide,  medulla  normal.  On  removal  of  capsule  the  surface 
is  found  very  irregular  and  elevations  from  the  surface  are  pale  red- 
gray-green  color.  These  elevations  do  not  correspond  with  any  change 
in  cortex  on  section.  Section  is  smooth,  solid,  markings  not  clear  but 
glomeruli  are  distineter  than  striae.  Line  between  cortex  and  medulla 
obscure.  The  left  kidney  contains  sand-like  calculi,  possibly  uratie, 
as  there  are  some  pale  areas  near  point  of  pyramid  which  are  firmer 
than  rest  of  tissue  and  rather  gritty.  Microscopic  section  of  kidney 
shows  the  architecture  much  disturbed  by  connective  tissue  overgrowth 
in  outer  layers  of  medulla  and  inner  layers  of  cortex  and  following  the 
medullary  rays  to  the  capsule.  Tubules  are  compressed  and  distorted 
in  the  vicinity  mentioned.  Epithelium  is  elsewhere  low,  opaque  and 
granular.  The  connective  tissue  about  the  glomeruli  is  thickened  and 
hyaline  as  it  is  in  most  other  places.  Tufts  are  not  yet  compressed. 
Following  medullary  rays  there  is  a  recent  round  and  polynuclear  in- 
filtrate both  around  and  in  tubules,    y 


284   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

A  Golden  Cat  {Felis  temminicki)  presented  sand  in 
the  urethra,  which  had  caused  a  traumatic  urethritis  and 
distention  of  the  bladder.  There  was  no  evidence  of  renal 
urate  collections,  but  a  low  grade  prostatitis  existed  so 
that  the  bladder  might  have  been  distended  before  the 
urethra  became  inflamed,  thereby  giving  opportunity  for 
sand  to  form  or  to  have  arisen  in  the  prostatic  ducts. 

'  Three  birds,  a  Bunting  (Passerina  ciris),  a  Bulbul 
{Chloropsis  aurifrons),  and  a  Buzzard  {Buteo  alhicau- 
datus),  had  large  cloacal  urate  calculi  which  could 
obstruct  the  ureter  but  had  failed  to  do  so;  one  had  an 
acute  ascending  pyelonephritis,  however.  / 

Excessive  urate  collections  in  ureters  and  kidneys 
occur  all  through  the  avian  orders  and  in  about  the  same 
percentages ;  meat  and  fish  eating  birds  have  practically 
no  cases,  however.  The  condition  seems  at  times  the  only 
finding  at  autopsy,  or  it  may  be  associated  with  uratic 
serositis.  Gout  of  birds  is  commonly  accompanied  by  it, 
but  need  not  be  since  two  of  the  best  examples  of  this 
disease  had  practically  noniial  kidneys. 

From  these  records  it  would  seem  that  renal  and 
pelvic  calculi  occur  almost  exclusively  in  herbivorous 
animals.  At  least  true  stones  forming  in  the  renal 
pyramid  and  pelvis  are  found  most  characteristically 
developed  in  the  Ungulata,  the  typically  herbivorous 
mammal.  Judging  by  the  bilateral  distribution  of  stones 
and  uratic  collections,  local  processes,  inflammation 
especially,  have  less  to  do  with  their  production  than  the 
availability  of  precipitable  inorganic  salts  in  the  urine. 

Tumors. 

Tumors  of  the  kidney  have  been  observed  sixteen  times, 
fourteen  of  which  were  primary  and  two  secondary.  The 
latter  two  concerned  an  epitheliomatous  metastasis  from 
a  malignant  papilloma  in  the  stomach  of  a  Kangaroo 
{Macropus  rufus)  and  a  sarcoma  growing  like  an  infarct 
secondary  to  a  mediastinal  tumor  in  a  Dorcas  Goat 
{Capra  hirciis).  The  only  important  primary  tumor  of 


Fig.  21. — CALCULI  FROM  RENAL  PELVIS  TO  END  OF  URETHRA.  COMMON 
RACCOON  (PROCYON  LOTOR).  THESE  STONES  WERE  PALE  YELLOW-GRAY.  THEY 
CONSISTED  OF  A  URATIC  BASE.  BUT  SOME  PHOSPHATES  AND  CARBONATES 
WERE  FOUND.  THE  RIGHT  KIDNEY  WAS  NOT  AFFECTED.  THERE  WAS  ONE 
IRREGULAR  CALCULUS  AND  THIRTY-SEVEN  SMOOTH  MASSES  FROM  BLADDER 
TO  END  OF  PENIS.  THIS  CASE  IS  NOT  INCLUDED  IN  STATISTICS.  OCCURRING 
AFTER  THEIR  COLLECTION   HAD   CEASED. 


THE  URINARY  TRACT  285 

the  kidney  in  a  mammal  was  found  in  a  Gray  Squirrel 
{Sciurus  carolinensis  pennsylv aniens),  a  solid  gray 
nodule  composed  histologically  of  large  and  small  deeply 
staining  cells,  many  containing  large  vacuoles  and  fitted 
with  a  small  dark  nucleus.  The  arrangement  of  the  ele- 
ments was  in  irregular  acini  or  bundles  and  thereby 
suggested  the  tumor  known  as  hypernephroma.  A  small 
nodular  adenoma  was  found  at  the  upper  end  of  the  right 
kidney  in  a  common  opossum  {Didelphys  virginiana) 
and  seemed  to  be  purely  of  renal  construction  in  that  an 
attempt  to  retain  tubular  arrangement  was  evident. 

Twelve  primary  tumors  occurred  in  birds,  and  of  these 
five  were  found  among  parrakeets,  they  being  curiously 
enough  all  of  the  same  type.  These  cases  were  all  dis- 
covered in  the  undulated  grass  variety  {Melopsiftacus 
undulatus  55666  )  and,  because  of  this  fact  and  their 
histological  similarity,  have  excited  interest.  Grossly 
they  are  irregularly  nodular  or  lobulated  tumors  usually 
springing  distinctly  from  one  lobe,  but  sometimes 
destroying  the  whole  organ;  they  are  soft,  resilient  and 
hold  their  place  well  during  manipulation.  Sometimes 
one  may  detect  the  topography  of  the  renal  lobes  on  cross 
section  while  at  other  times  the  mass  is  homogeneous. 
Microscopically  one  finds  the  structure  of  papillary 
adenoma  with  cystic  formations  or  the  production  of 
atypical  solid  nests  of  epithelia  which  would  have  to  be 
called  cancerous,  for  they  certainly  make  no  attempts  to 
retain  acinus  or  duct  groupings.  Carcinomatous  areas 
have  been  discovered  in  two  of  these  cases,  not  in  the 
other  three,  which  have  been  called  papillary  adenoma. 
One  of  the  tumors  was  subjected  to  many  sectionings  and 
different  stainings  techniques  to  discover,  if  possible, 
animal  and  vegetable  parasites ;  this  search  failed.  One 
of  these  tumors  produced  hemorrhage  by  rupture  of  a 
pyramid  but  extension  to  adjacent  tissue  and  metastases 
have  not  been  seen.  Pathologically  these  must  be  classi- 
fied with  the  tumors  but  because  of  the  number  of  closely 
similar  growths  in  the  same  avian  species  housed  in  the 


286   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

same  enclosure,  the  possibility  of  a  parasitic  cause  will 
not  be  forgotten. 

The  remaining  seven  renal  neoplasms  are  made  up  of 
two  adenomata  in  a  Jungle  Babbler,  {Crater opus  ccmo- 
rw5)j  and  a  red-headed  duck  [FuUigula  ferina  americana) , 
two  adenocarcinomata  in  a  saffron  finch  9  {Sy calls 
-fiaveola),  and  a  chestnut-eared  finch,  6  {Amadina  casta- 
notis),  two  hypernephromata  in  an  American  robin,  9 
(Planesticus  migratorius)  and  a  European  blackbird,  9 
(Merula  ynerula),  and  a  spindle-celled  sarcoma  on  a  scaly 
ground  dove  9  {Scardapella  squamosa).  One  adeno- 
carcinoma sent  out  metastases  to  the  lung,  one  hyper- 
nephroma had  secondarj^  growths  in  the  lungs,  the  other 
in  the  liver.  The  sarcoma  case  presented  a  metastasis 
in  the  tibia. 

Diseases  of  the  lower  parts  of  the  urinary  tract  are 
not  numerous  and  of  incidental  interest  only;  many  are 
associated  with  or  due  to  lesions  in  the  genital  organs,  and 
will  be  referred  to  later.  Cystitis  is  uncommon,  only 
being  observed  some  four  times  unrelated  to  prostatitis 
and  vesiculitis.  Two  of  these  cases  were  secondary  to  a 
traumatic  urethritis.  No  stones  have  been  seen.  A  mixed 
cell  sarcoma  was  f omid  springing  from  the  bladder  wall  in 
a  Richardson's  Kangaroo  Rat  {Perodipus  richardsoni). 
The  written  record  has  unfortunately  been  lost,  but  the 
preserved  slide  confirms  the  original  diagnosis.  Rupture 
of  the  bladder  occurred  in  a  Gray  Fox  with  stenosis  of  the 
end  of  the  penile  urethra  causing  retention  and  secondary 
cystitis.  Another  stricture  of  the  urethra  in  an  Ocelot 
{Felis  chibigonazon)  caused  great  dilatation  of  the  blad- 
der. This  animal  is  thought  to  have  chewed  off  all  the 
external  genitalia  because  of  lice,  with  the  result  that  the 
stump  of  the  urethra  became  involved  in  a  contracting 
cicatrix.  Opossums,  raccoons  and  wild  dogs  have  shown 
light  cases  of  urethritis  seemingly  traumatic  in  origin, 
and  two  dogs  had  cystitis  and  urethritis  associated  mth 
what  was  believed  to  be  distemper. 


SECTION  IX 
THE  FEMALE  REPRODUCTIVE  ORGANS 

The  mechanism  and  organs  of  reproduction  differ  so 
widely  in  the  classes  under  discussion  that  it  will  be 
necessary  to  describe  separately  the  alterations  in  mam- 
mals and  in  birds.  Examples  of  abnormality  and 
disease  are  not  very  numerous  and  I  shall  cite  cases  for 
many  of  the  conditions  rather  than  prepare  comparative 
lists  as  has  been  possible  in  many  foregoing  sections. 
Some  years  ago  Dr.  Edward  A.  Schumann  (1)  studied  the 
comparative  anatomy  and  physiology  of  the  mammalian 
female  generative  organs,  and  I  shall  condense  and  para- 
phrase his  work.  (The  complete  articles  may  be  found 
as  in  the  references  below.)  This  gentleman,  because  of 
his  gynecological  experience  and  broad  interest  in  com- 
parative biology,  has  been  consulted  whenever  unusual 
material  from  this  tract  has  presented  itself  so  that  many 
of  the  descriptions  that  I  shall  employ  are  due  to  him.  I 
take  this  opportunity  to  acknowledge  with  thanks  his 
interest  and  helpfulness. 

Comparative     Anatomy    in     Explanation     of     Human 

Uterine  Anomalies. 

The  development  of  the  genital  tract  seems  to  be 
essentially  the  same  through  all  orders  in  that  the  genital 
ridge  forms  the  ovary  and  its  attachments  while  the 
MuUerian  ducts  supply  the  tubes,  uterus  and  vagina. 
Early  in  fetal  life  these  two  longitudinal  ducts  begin  to 
approach  one  another,  and  by  the  end  of  the  third  month 
should  be  in  the  position  which  they  are  to  retain  for  the 
full  development  of  their  end  result.    In  the  human  being 

( 1 )  Comparative  Anatomy  of  the  Female  Genitalia,  Am.  Jour,  of 
Obstet.,  Vol.  LXIV,  No.  4,  1914.  Mechanism  of  Labor  from  a  Comparative 
Standpoint,  Ibid.,  Vol.  LXIX,  No.  4,  1914.  Dynamics  of  the  Female  Pelvis, 
its  Evolution,  etc..  Ibid.,  Vol.  LXXI,  No.  1,  1915. 

287 


288  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

this  position  is  complete  union  and  fusion,  with  the  pro- 
duction of  a  single  tube  from  the  uterine  fundus  to  the 
exterior,  while  in  the  lowest  mammals,  edentates  and 
marsupials,  the  two  Mullerian  ducts  retain  their  lateral 
position,  and  upon  completion  of  embryonal  life  a  double 
tube  from  the  ovaries  to  the  exterior  is  found.  If  the  nor- 
mal fusion  of  the  ducts  does  not  take  place,  and  if  for  any 
reason  their  proper  relation  is  not  reached,  an  abnormal- 
ity will  result  varying  according  to  the  stage  of 
development  that  has  been  reached.  Thus  in  man  instead 
of  a  single  uterus  of  triangular  shape  and  a  single  cervix, 
a  double  set  of  tubes  may  be  found.  It  can  be  shown  that 
the  abnormaUties  of  the  human  uterus  are  of  definite  and 
fixed  types  corresponding  to  an  arrest  of  fusion  or  com- 
pletion of  the  developmental  cycle  as  given  for  the 
Mullerian  ducts  and  further  that  these  very  deformities 
are  comparable  to  normal  organs  of  lower  orders.  In 
other  words  abnormalities  in  arrest  of  development  in 
the  human  uterus  represent  normal  types  of  lower  uteri 
at  various  evolutionary  stages.  In  monotremes  there  are 
two  ovaries,  tubes,  uteri,  cervices,  a  urogenital  passage 
and  a  clitoris.  This  corresponds  with  Uterus  didelphys 
with  a  single  vagina.  This  abnormality  while  occurring 
in  the  lowest  mammal,  does  not  represent  the  most 
marked  deformity  known  for  the  human  being — that  in 
which  the  double  tube  remains  to  the  vaginal  outlet,  a 
condition  found  in  the  marsupials.  In  tliis  order  the 
uteri  are  entirely  separate,  and  each  is  fitted  with 
its  own  vagina.  The  next  higher  order,  Edentata,  seems 
somewhat  out  of  place  if  it  be  judged  by  its  female 
genitalia  since  it  is  possessed  of  a  triangular  uterus 
and  single  vagina  but  without  distinct  cervical  seg- 
ment ;  the  tract  is  very  similar  to  that  of  the  Primates. 

Rodents,  in  the  various  families,  present  no  uniform 
uterine  construction,  there  being  four  kinds  corre- 
sponding to  as  many  degrees  of  Mullerian  duct  fusion 
and   differentiation.      The   lowest    forms    simulate    the 


THE  FEMALE  REPRODUCTIVE  ORGANS  289 

marsupials,  another  group  is  like  the  monotremes,  a 
third  shows  a  complete  fusion  with  a  single  cordiform 
uterine  body,  while  the  last  resembles  somewhat  the 
first,  but  the  uterine  di\dsions  are  bound  together  and 
the  vagina  is  divided  only  half  way  do^vn.  The  Insec- 
tivora  occupy  a  transitional  position  resembling  higher 
and  lower  groups  in  having  a  long  uterovaginal  canal, 
without  distinct  cervix,  extending  upward  into  long 
curved  comua.  Cetacea  (whales)  have  a  highly  rugous 
single  vagina,  a  distinct,  short  uterine  segment  divided 
into  two  separate  horns.  The  genitalia  of  Sirenia 
resemble  those  of  the  last  group,  but  the  cervix  is  better 
developed.  In  the  last  three  orders  the  clitoris  begins 
to  be  well  developed  and  to  present  externally.  Probos- 
cidea  have  a  single  vagina  separated  from  the  short 
uterine  body  by  three  transverse  folds  corresponding  to 
the  cervix;  the  two  cornua  are  long  and  wide. 

''The  foregoing  orders  present  in  their  uteri  all  the 
essential  characteristics  of  uterus  bicomis  unicollis  with 
single  vagina  and  are  therefore  the  homologues  of  this 
anomaly  in  man." 

In  the  Perissodactyla,  the  bicornate  uterus  has  a 
body  of  a  little  less  than  half  the  whole  length;  there  is 
a  sphincter  at  the  lower  end  of  the  body  but  no  project- 
ing cervix.  The  cornua  are  longer  still  in  the 
Artiodactyla  and  are  coiled  in  a  manner  suggesting 
spiral  sheep's  horns;  there  is  a  differentiated  projecting 
cervix.  In  cats  the  length  of  the  uterine  body  and  of  the 
cornua  are  almost  equal  and  both  are  flat  tubes;  the 
cervix  is  prominent  and)  the  vagina  long  and  rather 
smooth.  The  dog's  uterus  is  similar  but  the  two  cornua 
are  bound  together  or  fused  before  the  point  at  which 
their  termini  enter  that  of  the  uterine  body;  the  cervix 
is  not  very  prominent,  but  well  formed  and  the  vagina 
is  rugous.  These  types  correspond  to  the  uterus  cordi- 
formis.  Lemurs  have  a  common  uterovaginal  cavity 
like  the  Edentata.  The  lower  monkeys  possess  a  long 


290   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

slender  uterus  with  definite  superior  lateral  angles,  the 
last  vestiges  of  the  cornua,  a  prominent  cervix  and  a 
short  rugous  vagina.  The  higher  apes  have  a  uterine 
construction  almost  identical  mth  that  of  man  at  the 
stage  of  infantile  development. 

**From  the  foregoing  study  it  is  clearly  shown  that 
every  anomaly  of  the  female  genitalia  in  Man  is  in 
reality  the  result  of  atavism  and  hence,  a  degenerative 
change,  and  inasmuch  as  every  special  form  of  anomaly 
finds  its  counterpart  in  the  normal  anatomical  arrange- 
ment of  the  analogous  structures  in  one  or  another  of 
the  great  mammalian  groups,  one  additional  item  of 
proof  is  offered  in  support  of  that  greatest  of  biological 
doctrines,  the  descent  of  Man  from  the  lower  forms 
of  life." 

Anatomy  of  Labor. 

In  a  second  article  Schumann  reviews  the  com- 
parative anatomy  of  labor,  demonstrating  that  the  basic 
principles  are  essentially  the  same,  alterations  only 
being  in  the  direction  of  accommodation  to  the  pelvic 
construction.  In  order  to  make  the  analysis  comparable 
with  human  conditions  the  pelvis  is  studied  as  if  the 
animal  were  standing  erect  upon  the  hind  hmbs. 

The  salient  points  of  difference  between  the  quad- 
ruped pelvis  and  the  biped,  human  type  may  be 
epitomized  as  follows:  (a)  The  entire  pelvis  lies  (with 
the  animal  in  its  normal  station)  in  a  generally  horizon- 
tal position  with,  a   slight  slope  downward  anteriorly, 

(b)  The  false  pelvis  is  almost  entirely  wanting,  there 
being  practically  no  bony  structures  above  the  brim  with 
the  exception  of  the  small  upper  portions  of  the  ilia. 

(c)  The  pelvis  in  quadrupeds  is  never  basin-shaped,  the 
lateral  walls  from  the  iliac  crests  to  the  tuberosities  of 
the  ischia  lying  roughly  parallel  to  each  other  and 
enclosing  a  pelvic  cavity  rectangular  in  outline.  Only 
in  the  highest  apes  does  the  basin-shaped  pelvis  appear. 


THE  FEMALE  REPRODUCTIVE  ORGANS     291 

(d)  The  angle  of  the  axis  from  the  promontory  of  the 
sacrmn  to  the  sjTuphysis  is  always  greater  in  the  quad- 
rupeds than  in  man,  averaging  in  the  former  from  70  to 
80  degrees,  in  the  latter  about  55  degrees,  (e)  The 
symphysis  pubis  is  an  extremely  long  joint,  being  fre- 
quently greater  than  half  the  length  of  the  entire  pelvis. 
(f)  The  sacroiliac  joint  is  more  or  less  movable  in.  all 
quadrupeds,  especially  in  young  animals.  The  rotation 
of  the  sacrum  on  the  ilia  increases  the  anteroposterior 
(dorsoventral)  diameter  of  the  outlet  and  at  the  same 
time  wedges  apart  the  ilia,  thus  increasing  the  lat- 
eral diameter. 

In  regard  to  the  forces  of  labor  it  is  to  be  pointed  out 
that  in  the  lower  mammals  the  pregnant  uterus  hangs  be- 
low the  pubic  arch  so  that  the  fetus  must  rise  at  an  angle 
of  about  45  degrees  to  pass  over  the  pelvic  brim.  In  so 
doing  it  meets  the  narrowest  part  of  the  triangular  bony 
pelvic  inlet,  the  anterior  pubic  angle.  Since  the  sacrum 
is  above  and  out  of  the  way,  the  lateral  diameter  is  the 
one  which  must  be  suitable  to  the  passage  of  the  pre- 
senting part.  This  is  the  head  in  homo,  the  largest  part 
of  the  fetus,  but  in  lower  animals  either  head  or  breech 
often  accompanied  by  one  or  more  extremities,  may 
present;  the  head  is  not  the  largest  part  in  lower  mam- 
malian fetuses.  The  uterine  contraction  proceeds  as  in 
man,  the  fundus  and  cornua  acting  alone  until  the  cervix 
contains  the  fetus,  at  which  time  all  parts  contract.  In 
the  bicornate  organ  both  sides  must  contract  or  the  fetus 
might  be  forced  from  the  gravid  to  the  empty  side.  In 
muciparous  animals  with  both  uterine  horns  occupied, 
the  fetuses  lie  head  to  head,  breech  to  breech  and  are 
expelled  alternately  from  each  side. 

In  uniparous  animals  rotation  is  in  the  nature  of  an 
accommodation  of  the  greatest  diameter  of  the  fetal 
body  in  cross  section  to  greater  axis  of  the  mother — the 
dorsoventral.  Uterine  contractions  cause  the  fetus  to 
unfold    from    its    elliptical    form    and    to    assume    an 


292   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

extended  position,  permitting  head  or  breech  to  enter 
the  lower  pelvis.  The  pubic  angle  ha\^ng  been  passed 
and  the  extension  of  the  presenting  part  being  success- 
fully accomplished,  there  is  no  striking  difference  in  the 
manner  of  external  expulsion  of  the  fetus. 

Dystocia. 

Dystocia  in  domesticated  animals  is  a  well  studied 
subject,  and  its  general  clinical  phases  are  fairly  Avell 
known.  Several  cases  have  been  observed,  details  of 
which  are  worthy  of  note  since  some  of  them  are 
entirely  complete. 

Inertia  uteri  as  a  single  non-obstructive  condition 
seems  not  to  have  been  observed  here,  although  well 
enough  kno^vn  to  veterinarians.  Nor  has  a  case  of 
dystocia  been  seen  as  the  result  of  excessive  expulsive 
force.  Obstruction  to  natural  passage  by  bony 
deformities  or  malformations  is  a  common  occurrence  in 
man  but  not  so  in  lower  animals.  There  may  also  be 
dystocia  by  reason  of  a  normally  formed  but  too  small 
pelvis  when  the  female  has  been  impregnated  by  a  much 
larger  male,  or  if  the  female  conceived  before  the  pelvis 
has  achieved  its  full  growth. 

Deformities  may  be  due  to  irregularities  of  bony 
development,  exostoses,  fracture  or  diseases  of  the 
osseous  system.  A  most  interesting  case  of  this  type 
occurred  in  a  Barbary  ape  {Macacus  innuus)  which  had 
been  in  the  collection  for  two  years  and  was  apparently 
in  good  health  when  discovered  in  labor. 

No  progress  being  made  and  the  animal  becoming  shocked,  an  ineffec- 
tual attempt  was  made  to  deliver  by  version,  the  monkey  dying  during 
the  operation.  Upon  autopsy  the  uterus  contained  a  fetus  apparently 
at  term.  The  head  was  extended  so  that  the  face  presented,  but  the 
head  was  not  engaged.  The  cervix  was  fully  dilated  but  the  uterine 
muscle  was  relaxed  and  flaccid.  The  fetus  Avas  dead  Avhen  the  animal 
was  first  examined.  The  uterus  contained  two  placenta?  as  is  normal 
for  these  apes,  the  left  placenta  being  the  place  of  attachment  of  the 
fetus  tvhile  the  right  one  was  somewhat  smaller  and  presented  no  umbili- 


THE  FEMALE  REPRODUCTIVE  ORGANS     293 

cal  cord.  The  fetus  was  normal  in  size  and  form,  the  face  was  extended 
and  its  lower  portion  far  advanced  in  a  caput  succedaneum.  The  meas- 
urements of  the  fetal  were  as  follows :  bitemporal  5.5  cm.,  biparietal  6 
em.,  occipitomental  8.5  cm.,  occipitofrontal  7.5  em.  The  pelvis  (dried  spec- 
imen) presents  a  most  interesting  condition.  The  sacrum  is  bent  sharply 
forward,  carrying  with  it  the  border  of  the  ilia,  which  are  bent  upon 
themselves  forward  and  downward.  The  lateral  walls  of  the  pelvis  are 
greatly  narrowed,  the  ischia  drawn  inward.  The  pubes  and  the  symphy- 
sis are  fairly  normal.  The  pelvic  measurements  are:  diagonal  con- 
jugate 6  cm.,  true  conjugate  4  cm.,  greatest  transverse  4  cm. 

It  is  apparent  at  a  glance  that  here  was  an  impossible 
labor,  since  the  head  of  the  fetus  could  not  possibly 
enter  the  pelvis,  the  size  of  which,  intra  vitam,  must  have 
been  less  than  the  above  measurements  by  reason  of  the 
soft  parts.  ''This  is  in  the  experience  of  the  writer  a 
unique  case  of  a  complete  obstetric  history,  plus  the 
specimens,  of  labor  with  an  osteomalacic  pelvis  in  a 
wild  animal." 

Another  case  may  be  added  to  those  already  reported 
by  Dr.  Schumann,  as  follows: 

Hairy  rumped  Agouti  {Dasyprocta  prymnolopha) .  Dystocia.  An 
apparently  normal  fetus  occupies  the  left  uterine  horn.  The  nose  was 
engaged  in  the  pelvis  and  has  been  moulded  in  a  curve  pointing  to  the 
right.  The  fetus  measures — bitemporal  30  mm.,  cervical-coronal  31 
mm.,  length  of  fetus  17  cm.  Pelvic  inlet  in  the  fresh  state  measures  18 
mm,  transversely  and  about  17  mm.  anteroposteriorly.  The  umbilical  cord  is 
10  em.  long  and  appears  normal  as  do  the  membranes.  Placenta  pre- 
sents as  a  spherical  mass  of  hard  dense  consistency,  35  mm.  in  diameter 
and  with  apparently  normal  placental  tissue  occupying  the  lower  border 
of  this  spherical  mass.  On  section  the  mass  shows  areas  of  alternating 
soft  red  tissue  separated  by  communicating  trabeculae  of  dense  white 
fibroid  tissue.  This  mass  is  distinctly  encapsulated,  but  the  nature  of 
the  enclosing  membrane  is  indeterminate.  The  pelvis  in  dry  state  shows 
evidences  of  malformation  due  to  trauma.  The  right  ileum  is  pushed 
forward  and  inward  carrying  the  acetabulum  a  short  distance  inward 
and  backward.  There  is  a  marked  thickening  about  the  right  acetabu- 
lum. At  the  upper  portion  of  the  symphysis  there  is  marked  bending 
backward  toward  the  sacrum  with  thickening  of  the  bone.  The  last 
sacral  vertebra  is  sharply  bent  and  anchylosed,  forming  an  angle  of 
sixty  degrees.  The  pelvic  measurements  in  the  dry  state — at  superior 
strait-transverse  24  mm.,  right  oblique  23  mm.,  left  oblique  26  mm.,  true 
conjugate  22  mm.,  outlet  17  mm.,  between  the  ischiatic  spines. 


294  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

This  is  obviously  a  traumatic  malformation  and 
forms  a  relatively  contracted  pelvis.  The  delivery  of  an 
adult  fetus  is  impossible. 

Cape  Hyrax  {Hyrax  capensis).  Impossible  labor  clue  to  malforma- 
tion of  the  pelvis.  This  animal  died  as  the  result  of  shock  and  exhaus- 
tion of  labor.  She  was  pregnant  of  two  fetuses,  one  of  which  was 
extracted  manually  by  the  keeper  but  was  dead  at  birth.  Twenty-four 
hours  later  the  animal  was  found  dead.  On  autopsy  there  was  present 
a  fully  developed  fetus  in  the  right  cornu,  the  head  just  above  the  pelvic 
inlet.  The  left  cornu  was  large  and  boggy.  The  myometrium  of  the 
right  side  was  so  thin  as  to  be  almost  transparent.  On  examining  the 
bony  pelvis  the  reason  for  the  dystocia  is  at  once  apparent.  The  sacrum 
is  tilted  to  the  right,  and  the  body  of  the  left  ileum  is  bent  sharply  to 
the  right,  the  pelvic  inlet  being  obliquely  contracted,  the  right  oblique 
diameter  being  16  mm.  while  the  left  is  23  mm. 

The  pelvic  obliquity  made  the  birth  of  a  full  sized 
fetus  impossible,  the  one  delivered  being  under 
developed.  The  myometrium  was  evidently  stretched 
almost  to  the  point  of  rupture  when  death  occurred. 

Obstruction  to  the  birth  canal  by  abnormalities  in 
the  soft  parts,  such  as  muscular  rigidity,  edema,  or 
inflammation,  tumors,  atresia  or  developmental  defects 
are  occasionally  seen  by  veterinarians  but  have  not  been 
encountered  here. 

Dystocia  from  uterine  displacements  are  uncommon 
except  such  as  may  depend  upon  the  failure  of  support 
by  the  abdominal  wall,  since  this  is  the  principal  support 
of  the  organ  especially  when  gravid.  Hernia  or  hys- 
terocele  is  known  and  anteflexion  has  been  seen., 
Torsion,  a  rare  human  condition,  is  not  uncommon  in 
lower  animals  probably  due  to  the  loose  dependent 
position  of  the  pregnant  cornua,  attached  only  to  the 
pelvic  walls  by  slender  inactive  suspensory  ligaments. 
A  case  may  be  reported,  not  originally  described. 

Canada  Porcupine  {Erethizon  dorsatus  dorsatus) .  Obstructed  labor. 
This  animal  died  from  exhaustion  due  to  an  impossible  labor.  On 
autopsy  the  abdomen  contained  a  moderate  amount  of  clear  serous 
fluid  with  one  small  blood  clot.  The  right  uterine  cornu  was  distended 
with  a  fetus  to  about  the  same  diameter  as  the  uterine  bodv.    This  cornu 


THE  FEMALE  REPRODUCTIVE  ORGANS     295 

was  congested  and  edematous  and  its  walls  very  thin.  The  cornu  was 
twisted  one  half  full  turn  from  right  to  left,  so  that  the  pregnant  por- 
tion overlaid  the  uterine  body  anteriorly.  The  torsion  produced  a  com- 
pression of  the  vessels  on  the  right  side  to  the  point  of  violent 
congestion  of  the  cornu.  There  was  no  apparent  rupture  of  the  uterine 
walls,  death  having  occurred  from  exhaustion.  The  fetus  and  its  mem- 
branes were  normal. 

Many  forms  of  abnormalities  in  position  are  recog- 
nized for  domesticated  animals,  but  since  we  know  so 
little  of  the  early  stage  in  the  wild  specimens  no  data 
can  be  given. 

Complicated  labor  in  lower  varieties  of  animals  will 
follow  lines  similar  to  those  for  man  and  domesticated 
animals.  Hemorrhage  from  trauma  is  not  common  at 
term,  but  several  cases  of  abortion  following  injury  have 
been  seen.  Postpartum  hemorrhage  might  be  expected 
in  the  higher  apes  which  have  a  large  discoid  placenta 
similar  to  the  human  form,  but  when  the  placenta  is 
more  loosely  attached  and  is  subdivided  as  in  lower 
forms,  such  bleeding  is  of  no  danger ;  when  a  cotyledon 
is  torn  from  the  ungulate  uterine  wall,  a  free  hemorrhage 
sometimes  occurs. 

Geoifroy's  Marmoset  (Leontocebus  geoffroyi).  Puerperal  relaxation 
of  the  utenis  with  fatal  hemorrhage.  The  uterus  is  4  cm.  long,  15  mm. 
wide  at  intertubal  line.  Uterine  wall  averages  2  mm.  in  thickness.  Perito- 
neal surface  is  smooth,  glistening  and  intact.  Uterine  musculature  is  soft 
and  relaxed.  Entire  organ  is  intensely  congested  and  on  section  uterine 
cavity  contains  a  large  firm  blood  clot  completely  filling  it.  Mucosa 
is  of  deep  purple  color,  shows  many  fragments  of  decidua  and  is  the 
seat  of  profuse  hemoiThage. 

Placenta  previa  is  very  rare.  A  row  of  cotyledons 
may  form  near  the  internal  os,  therefore  like  a  placenta 
previa,  but  it  appears  to  be  of  no  consequence.  Pre- 
mature separation  of  the  placenta  is  known  to 
veterinarians,  and  is  exemplified  by  the  following  case 
in  our  records : 

Black  Lemur  (Lemur  macaco)  was  found  dead  in  its  cage.  Upon 
autopsy  the  uterus  contained  a  small  fetus  with  one  leg  and  the  tail 


296  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

protruding  from  the  vulva.  The  fetus  was  normal  in  size.  There  was 
a  large  amount  of  free  blood  in  the  uterine  cavity  and  extensive  extra- 
vasation into  the  myometrium.     The  placenta  was  completely  detached. 

Rupture  of  the  uterus  and  cervix  have  not  been  seen, 
but  a  traumatism  of  the  vagina  gave  occasion  for  the 
following  death; 

Bactrian  Camel  {Camelus  bactrianus)  died  of  shock  in  labor.  Upon 
autopsy  the  animal  was  found  to  have  hydatid  disease  of  the  liver,  lungs, 
and  spleen,  cirrhosis  of  the  liver,  and  nephrolithiasis.  Protruding  from 
the  vulva  was  a  portion  of  the  fetal  membranes,  the  whole  vaginal  wall 
and  several  coils  of  intestine  which  had  escaped  through  a  large  rent 
in  the  posterior  vaginal  wall.  The  anterior  wall  was  swollen  and  edema- 
tous, the  whole  region  surrounded  by  clotted  blood.  The  cervix  was 
obliterated,  the  membranes  unruptured,  the  fetus  in  the  normal  extended 
head  presentation.  The  veil-like  placenta  was  somewhat  injected  but 
otherwise  normal.  There  Avas  some  hemorrhage  about  the  rectum  but 
none  in  the  free  peritoneum. 

This  animal  had  broken  her  hind  leg  just  above  the 
fetlock  three  weeks  before  falling  into  labor,  and  was 
unable  to  stand.  It  is  evident  that  the  difficulty  of 
delivery  associated  with  an  unnatural  and  forced 
posture  due  to  the  fractured  leg  was  sufficient  cause  for 
the  rupture  of  the  vagina  where  the  tissues  were  degen- 
erated as  a  result  of  the  coincident  general  disease. 

Inversion  of  the  uterus  is  one  of  the  common  acci- 
dents of  labor  among  all  animals,  most  frequently  seen 
in  ruminants.  It  is  a  condition  more  to  be  expected  in 
lower  animals  than  in  man  because  of  the  long  slender 
relaxed  suspensory  ligaments,  the  length  of  the  uterus 
and  the  rigor  of  the  contractions.  The  immediate  causes 
are  those  operative  for  human  beings.  Three  cases  are 
recorded — one  in  an  axis  deer,  one  in  an  opossum,  one 
in  a  mouse,  the  last  being  detailed  in  the  following  notes : 

A  Japanese  Waltzing  Mouse  (Mus  wagnerii  rotans)  died  a  few 
hours  after  an  uneventful  labor.  On  autopsy  the  entire  uterus  was 
found  inverted  and  prolapsed,  the  organ  the  seat  of  a  violent  conges- 
tion, the  animal  having  died  of  shock. 


Fig.  22. — INVERTED  AND  PROLAPSED  UTERUS. 
JAPANESE  WALTZING  MOUSE  (MUS  WAGNERII 
ROTANS).      UTERUS  SHOWN  LYING   ON   CARD. 


THE  FEMALE  REPRODUCTIVE  ORGANS     297 

The  Pelvis. 

A  study  of  the  dynamics  of  the  female  pelvis  from  an 
evolutionary  standpoint  may  explain  some  of  the  diffi- 
culties attending  parturition.  Starting  from  the  biologi- 
cal law  that  morphology  follows  function  and  that  the 
anatomy  of  a  part  alters  to  suit  a  changed  physiology 
with  such  modifications  as  are  necessary  to  fit  each  part 
properly  to  interact  with  other  structures  comprising 
the  entire  animal,  it  is  evident  that  two  great  changes 
have  occurred  in  the  evolution  of  homo — the  assumption 
of  the  upright  posture  and  an  increase  of  intellectual 
power  necessitating  a  larger  cranium  of  modified  form. 
To  this  end  also  the  pelvis  would  have  to  change  both 
for  support  and  to  allow  the  passage  of  the  enlarged 
head.  The  functions  of  the  pelvis  are  (1)  to  attach  the 
legs  or  hinder  limbs  to  the  trunk;  (2)  to  furnish  points 
of  attachment  and  fulcra  for  the  great  muscles  which 
move  the  limbs,  and  in  the  case  of  man,  hold  the  trunk 
erect;  (3)  to  provide  egress  and  support  for  the  termi- 
nal canals  of  the  intestinal  and  urinary  systems ;  (4)  to 
provide  for  a  birth  passage;  (5)  to  act  as  a  shelf  and 
support  for  the  abdominal  viscera.  In  quadrupeds  the 
first  four  functions  being  perfectly  served,  there  are 
fewer  abnormalities  of  reproduction  (and  in  the  posi- 
tions of  viscera  as  well — Ed.).  In  man  natural  selection 
weeded  out  narrow  pelves,  but  the  present  product  is  as 
yet  an  imperfect  structure  for  one  of  its  main  uses, 
parturition.  It  should  be  a  funnel-shaped  basin  of  the 
shape  of  the  fetal  head  and  of  the  same  height  at  all 
points — that  is  not  oblique,  there  should  be  no  promon- 
tory, the  pelvic  symphysis  should  be  short  and  the 
sacrum  of  the  same  height.  This  would  obviate  internal 
rotation  now  necessitated  by  the  oblique  pelvis ;  this  does 
not  occur  in  quadrupeds. 

Tracing  the  evolution  of  the  pelvis,  it  is  to  be  found 
first  in  fishes  where  it  is  a  loose  disjointed,  variable 
structure  not  attached  to  the  spine;  in  some  it  consists 

20 


298   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

merely  of  iscliia.  In  reptiles  the  box  assumes  a  form 
suggesting  higher  types.  Passing  from  the  toads  to 
turtles  and  to  the  crocodiles,  the  elements  which  go  to 
make  up  the  pelvis  assume  a  more  and  more  osseous 
character  and  become  more  and  more  definitely  articu- 
lated mth  the  spine  or  with  the  differentiated  sacrum. 
Its  purpose  in  these  low  forms  is  mostly  as  a  support 
for  the  muscles  of  the  legs  and  back.  Birds  present  an 
advance  in  pelvic  construction  but  with  a  great  prepon- 
derance of  the  vertebral  column  since  thirteen  to 
seventeen  bones  may  fuse  to  form  a  sacral  "roof."  The 
ilia,  ischia  and  pubis  are  firmly  combined.  The  box  is 
long  and  narrow  with  a  heavy  part  for  the  acetabulum 
and  broad  surfaces  for  muscular  insertions.  The  pelvis 
of  birds  differs  from  that  of  cold-blooded  vertebrates 
in  the  greater  number  of  vertebral  segments  entering 
into  its  composition,  and  in  their  bony  confluence.  It 
differs  from  that  of  mammals  by  being  unclosed  by  an 
anterior  s\Tiiphysis  and  by  a  widely  perforate  acetab- 
ulum. The  ossification  of  the  pelvic  bones  is  to  afford 
a  support  for  the  legs,  and  the  open  pelvis  allows  passage 
of  the  large  brittle  egg.  The  shape  of  the  pelvis  is  of 
little  importance  in  parturition  in  the  foregoing  animals, 
except  for  birds  which  bear  large  eggs  when  the  pubis 
anteriorly  is  open  for  that  purpose. 

In  monotremes  one  finds  the  reptilian  type  of  pelvis 
with  the  three  divisions  of  the  innominate  bone  remain- 
ing separate.  The  pelvis  is  short,  hea^'y  and  flat  and  is 
fitted  with  marsupial  bones.  The  marsupial  pelvis, 
possessing  parallel  walls  made  by  the  ilia  and  ischia 
and  a  long  symphysis,  resembles  roughly  a  triangular 
prism.  The  sacrum  is  wedge-shaped,  mthout  a  promon- 
tory and  has  a  considerable  movability.  The  marsupial 
bones  are  quite  long.  The  triangular  outlet  is  many 
times  the  size  necessary  for  the  passage  of  the  fetus. 

The  rodent  pelvis  is  difficult  to  summarize  morpho- 
logically  by    reason    of    the    great    diversity    of    form 


THE  FEMALE  REPRODUCTIVE  ORGANS     299 

occurring  in  the  many  genera  of  the  order,  but  it  may  be 
said  that  here  the  pelvis  is  usually  of  a  type  rather 
higher  in  the  scale  than  the  other  structural  character- 
istics of  the  order  would  indicate.  The  outlet  is  more 
commonly  ovate  than  rectangular,  the  ischia  and  ilia 
lie  at  a  more  marked  angle  to  each  other,  and  the  true 
conjugate  forms  a  lesser  angle  than  is  common  in  quad- 
rupeds. The  sacroiliac  joint  is,  in  general,  freely  movable. 

The  female  insectivorous  pelvis  is  relatively  large, 
the  sacroiliac  junction  usually  being  long  and  well  knit 
while  the  pubes  are  slender,  wide  of  angle  as  to  their 
descending  rami  and  there  may  be  no  symphysis,  as  in 
bats.  Edentates  have  distinct  bony  unions  of  the  ele- 
ments of  the  innominate  bone  and  of  this  to  the  sacrum 
which  increases  in  width  downward  permitting  a  long 
synostosis  with  ischia  and  pubes;  these  joints  are  not 
movable.  The  pubes  are  slender  and  the  symphysis 
short.  The  sacrosymphyseal  angle  is  80  degrees  in  the 
armadillo.  The  inlet  is  roughly  triangular  to  almost 
round.  Cetacea  have  no  clearly  developed  pelvis,  its 
place  being  represented  by  two  long  bones,  larger  in 
males,  which  seem  to  be  the  insertion  of  the  genital 
erector  muscles.  There  is  no  junction  to  form  a  pelvis 
nor  is  there  an  acetabulum.  A  pelvic  box  is  absent  in 
the  Sirenia,  but  lateral  processes  from  the  lumbar  verte- 
brae form  a  sort  of  ileum  between  which  an  ischium 
is  located. 

Proboscidea  have  a  massive  pelvis  lying  vertical  to 
the  spine.  The  iliac  alae  are  wide  and  deeply  concave; 
the  ischia  are  short,  heavy  and  parallel  to  the  ilia;  the 
pubes  are  short  and  combined  in  a  heavy  symphysis ;  the 
sacroiliac  joint  is  short,  heavy  and  slightly  movable; 
the  outlet  is  ovate.  In  one  specimen  examined  the  length 
was  4  feet,  symphysis  18  inches,  true  conjugate  19 
inches,  transverse  diameter  17.5  inches;  crests  of  ihac 
were  28  inches  long. 


300   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Perissodactylic  animals  throughout  this  suborder 
have  similar  pelves  except  in  so  far  as  the  obliquity  is 
concerned.  In  the  rhinoceros  it  is  90°  with  the  spinal 
column,  in  the  horse  135° ;  the  angle  of  the  true  conju- 
gate is  10°  in  the  former,  50°  in  the  latter.  The  lateral 
halves  of  the  box  are  heavy  and  parallel ;  the  symphysis 
is  short  in  the  rhinoceros,  large  in  the  horse ;  the  outlet 
is  ovoid.  Artiodactyla,  including  pachyderms  and  rumi- 
nants, show  a  variety  of  shapes  and  constructions. 
Hippopotami  have  a  short  massive  box  tilted  at  about 
150°  from  the  spine,  with  widely  flaring  ilia  and  ischia; 
the  pubes  are  slender  but  not  combined  in  a  strong  sym- 
physis. In  ruminants  the  sacrum  consists  of  four  fused 
units ;  the  sacroiliac  joint  is  fairly  movable ;  the  obliquity 
is  about  145°;  ilia  are  long  and  slender  and  flaring;  the 
ischia  are  broad  and  parallel  with  the  ilia;  pubes  are 
slender  but  form  a  symphysis  about  two-fifths  the  total 
pelvic  length ;  the  inf rapelvic  angle  is  mde ;  the  outlet  of 
the  pelvis  is  almost  rectangular. 

The  general  characteristics  of  the  carnivorous  pelvis 
may  be  summarized  as  consisting  of  a  long  strong 
symphysis,  parallel  lateral  pelvic  walls,  a  great  sacro- 
symphyseal  angle,  and  a  marked  separation  of  the  bodies 
of  the  ischia.  The  sacroiliac  joint  is  in  general  mod- 
erately movable. 

The  pelvis  of  lemurs  is  narrow,  attached  lightly  to 
the  slender  sacrum,  tilted  at  an  angle  of  140°  and  is 
possessed  of  a  short  weak  symphysis;  it  resembles  the 
structure  in  bats.  In  macaques  the  box  is  long,  the 
sacrum  wide,  with  a  short  iliac  synostosis,  the  ilia  long, 
narrow  and  curved  out  sharply,  the  ischia  are  continuous 
with  the  ilia  and  widely  separated;  the  symphysis  is 
short,  about  one-sixth  the  pelvic  length;  the  angle  of 
the  superior  strait  is  about  60° ;  the  outlet  is  oval,  the 
transverse  diameter  being  short.  The  chimpanzee  pelvis 
is  made  up  of  a  wedge-shaped  sacrum  composed  of  three 
vertebrae,  wide,  flaring,  concave  ilia,  stout  well  separated 


THE  FEMALE  REPRODUCTIVE  ORGANS     301 

ischia  with  flattened  tuberosities  and  a  short  symphysis 
parallel  to  the  sacrum ;  the  outlet  is  ovoid ;  true  conjugate 
angle  is  65°;  the  sacroiliac  junction  has  little  motility. 
In  the  gorillas  the  following  points  differ  from  the  last 
described  structure.  Five  vertebrae  comprise  the  sacrum 
and  the  anterior  surface  is  distinctly  concave ;  the  pelvic 
contour  while  still  ovoid,  has  the  two  diameters  more 
nearly  equal;  the  pelvic  angle  is  obtuse;  the  true  conju- 
gate is  at  an  angle  of  70°. 

In  man  the  salient  features  of  the  pelvis  are — a 
broad,  wedge-shaped  sacrum,  concave  anteriorly,  with 
wide  articular  surfaces  and  a  limited  motility;  widely 
flaring  ilia  including  the  concave  curvature  of  the  body 
of  the  bone  whereby  the  lateral  diameter  of  the  pelvic 
inlet  becomes  wider  than  in  lower  orders;  short  stout 
pubes  with  a  narrowed  angle  beneath  them ;  heavy  blunt 
ischia  with  large  tuberosities;  true  conjugate  is  at  an 
angle  of  55°.  The  human  fetal  pelvis  resembles  that 
of  quadrupeds. 

Study  of  these  data  indicates  that  the  quadruped 
pelvis  retains  many  things  in  common  through  all  the 
orders  especially  in  being  a  roughly  rectangular  struct- 
ure lying  chiefly  horizontally,  with  a  poorly  developed 
false  pelvis,  straight  ischia  and  a  long  symphysis  pubis ; 
the  angle  of  the  true  conjugate  is  greater  than  in  man  and 
may  be  up  to  80°.  The  long  pubic  synostosis  changes 
the  relation  of  the  true  and  diagonal  conjugate,  but  the 
former  is  no  indicator  of  pelvic  capacity,  since  in  lower 
mammals  the  promontory  of  the  sacrum  lies  anterior  to 
the  symphysis ;  the  vertical  diameter  is  a  better  measure 
of  pelvic  size  and  form.  Sacral  movement  seems 
greater  in  lower  animals  especially  in  youth.  The  shape 
of  the  pelvic  inlet  is  triangular  in  the  lowest  forms,  the 
posterior  base  of  this  becoming  wide  as  one  ascends  in  a 
zoological  line ;  the  concavity  of  the  ilia  also  increases  so 
that  the  higher  the  animal  the  more  curved  are  the  lat- 
eral   borders.      In     quadrupeds     the     anteroposterior 


302   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

diameter  is  greater  than  the  lateral;  the  reverse  is  true 
in  man.  The  long  straight-sided  quadruped  pelvis  is 
retained  because  of  the  direction  of  the  forces  from  the 
legs,  which  is  as  much  or  more  upon  the  anterior  or 
pubic  arch  as  upon  the  sacroiliac  junction.  In  the  semi- 
upright  position  of  the  monkey  the  force  is  directed 
backward  and  do^aiward  upon  the  sacrum,  this  aiming 
to  widen  the  pelvis  by  forcing  the  ilia  apart.  The  effect 
of  the  upward  force  from  the  femora  is  to  throw  the 
pelvis  upward  and  anteriorly  by  directing  the  line  of 
action  through  the  acetabula  more  toward  the  ventral 
surface.  The  sacroiliac  ligaments  hold  the  ilia  firmly, 
their  alae  being  spread  outward  by  the  force  from  above. 
Elevation  of  the  pubes  shortens  their  s\Tnphysis  and  the 
true  conjugate.  By  these  changes  the  birth  canal  is 
shorter,  entirely  bony,  with  the  upper  inlet  on  the  same 
plane,  and  promontory  and  symphysis  are  near  enough 
the  same  level  to  be  met  at  the  same  time  by  the  engaging 
head.  In  these  pelves  the  anteroposterior  diameter  is 
still  long  and  superior  rotation  is  not  necessary. 

In  Man  the  force  exerted  on  the  pelvis  from  above  is 
greatest  among  all  animals  and  is  greater  when  he  is  in 
active  motion.  The  force  is  directed  from  above  to  the 
sacroiliac  joint,  the  iliac  bodies  and  the  acetabular 
region  while  from  below  the  pressure  is  directly  exerted 
upon  the  last  named.  The  force  from  above  rotates  the 
sacrum  downward  at  its  upper  end,  the  attached  sacro- 
iliac ligaments  at  the  same  time  pulling  the  alae  inward 
and  throwing  outward  the  lower  end,  the  acetabular 
part.  This  tends  to  widen  the  pelvic  box  and  to  reduce 
its  anteroposterior  diameter.  But  the  force  acting  from 
the  legs  and  the  adductors  of  the  thigh  push  the  lower 
parts  of  the  ilia  and  the  ischia  and  pubes  apart,  thus 
counterbalancing  the  effort  of  the  force  from  above.  The 
combined  forces  tend  to  bend  the  iliac  bodies,  thereby 
producing  the  curved  lateral  margins  of  the  superior 


THE  FEMALE  REPRODUCTIVE  ORGANS     303 

strait  and  making  the  lateral  axis  long.  As  between 
these  two  forces  that  from  above  is  certainly  the  greater. 

The  effect  of  these  forces  can  be  followed  by  compar- 
ing a  quadruped  pelvis,  a  human  fetal  pelvis  and  an 
adult  human  one.  The  first  two  are  similar  chiefly  in 
the  shape  of  the  inlet  and  the  flat  character  of  the  ilia. 
This  is  strongly  corroborative  of  the  evolutionary 
development  of  the  pelvis  to  meet  the  demand  of  the 
upright  position.  The  effects  of  this  evolution  are  as 
follows :  First,  to  develop  a  forward  inclination  of  the 
sacrum  and  a  concavity  in  its  surface  anteriorly,  second 
to  increase  greatly  the  iliopubic  and  ilioischiatic  angles, 
third  to  cause  the  acetabula  to  move  forward  of  the  lumbo- 
sacral axis,  fourth  to  shorten  the  bodies  of  the  ilia  and 
to  develop  in  them  a  regularly  curved  surface,  the  con- 
cavity of  which  faces  forward  and  inward,  fifth  to 
decrease  the  interpubic  angle  and  accordingly  to  remove 
the  triangular  quality  of  the  pelvic  contour,  sixth  to 
increase  the  transverse  diameter  at  the  expense  of  the 
anteroposterior,  and  seventh  to  decrease  greatly  the 
sacrosymphyseal  angle  with  the  result  that  the  entire 
pelvic  cavity  lies  in  one  plane. 

The  effect  of  forces  in  alterations  of  the  pelvic  archi- 
tecture may  be  seen  in  their  several  stages  by  the 
observation  of  the  mammalian  pelvis  in  the  course  of  its 
evolution  and  development.  The  transitions  in  form  are 
very  gradual,  but  their  gradations  are  well  shown  in  the 
characteristic  forms  which  have  been  described ;  the  quad- 
ruped, monkey,  anthropoid  ape,  human  fetal  and  adult 
human  type. 

By  an  examination  of  the  adult  pelvis  and  fetal  skulls 
it  will  be  seen  that  the  shape  of  the  quadruped  birth  canal 
accommodates  the  fetal  head  nicely  when  in  extension 
since  there  is  no  large  posterior  cranial  development,  the 
head  and  neck  being  of  nearly  the  same  thickness.  In  the 
monkey,  where  the  facial  angle  increases,  the  head  does 
not  advance  as  a  pointed  presentation  but  as  an  irregular 


304   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

surface — the  chin,  forehead  or  occiput.  It  is  only  when 
the  head  has  assumed  a  high  facial  angle  and  a  well 
developed  occipital  lobe  and  the  pelvic  inlet  has  become 
transversely  wide,  that  internal  rotation  is  necessary. 

Resuming  a  discussion  of  our  records  I  shall  at  this 
place  introduce  additional  cases  of  pathology  in  the 
parturient  genital  organ  and  then  discuss  the  general 
system.  Beside  the  cases  of  dystocia  discussed  in  pre- 
ceding pages  there  occurred  in  a  deer  a  complete  eversion 
of  the  uterus,  which  was  incompletely  involuted,  accom- 
panying prolapse  of  the  rectum  and  bladder  due  to 
straining  after  severe  injuries,  probably  inflicted  by  a 
male.  The  animal  lived  long  enough  to  develop  a  purulent 
cystitis  which  spread  to  the  vagina,  uterus  and  pelvic  soft 
parts.  The  left  uterine  cornu  of  an  American  bison  was 
found  to  contain  a  decomposing  fetus  at  about  half  term ; 
general  sepsis  had  supervened  which  led  to  the  animal's 
death  in  about  a  week,  according  to  the  keeper 's  observa- 
tions. The  history  of  a  monkey,  followed  through  two 
pregnancies  and  finally  dying  of  tuberculosis  and  sepsis 
is  interesting. 

Pigtailed  Macaque  [Macacus  nemestrinus) .  Received  March  5, 1903, 
died  October  29,  1904.  This  animal  gave  birth  on  December  28,  1903  to 
a  young  one  which  appeared  feet  first;  the  arms  appeared  to  have  pene- 
trated the  septum  between  the  vagina  and  rectum  and  protruded  through 
the  anus.  Had  to  be  cut  off  before  delivery.  She  had  a  second  young 
one  October  21,  1904,  which  was  properly  delivered  but  was  weak  and 
lived  two  days,  having  received  little  attention  from  the  mother.  The 
mother  since  has  been  weak  in  the  hind  legs,  eaten  almost  nothing  and 
several  times  has  passed  a  little  blood  in  the  stool.  Lungs  are  partially 
collapsed;  contain  scattered  small  pinhead  tubercles;  right  lower  lobe 
contains  a  caseous  nodule  about  the  size  of  a  cherry.  Both  pleurjB  con- 
tain clear  fluid.  Abdomen  contains  a  half  pint  of  cloudy  fluid.  Omen- 
tum adherent  along  lower  border  and  region  of  spleen  and  contains 
pinhead  tubercles.  Surface  of  the  liver  is  studded  with  numerous  pin- 
point, pinhead  and  (old)  pea  sized  tubercles.  Some  places  in  the  liver 
seem  to  be  the  result  of  two  invasions.  Spleen  presents  small  cherry 
sized  tubercles  thickly  grouped  together  leaving  small  amount  of  splenic 
tissue  to  be  seen.  Kidneys  contain  pinhead  to  barley  grain  tubercles. 
Uterus  is  somewhat  enlarged.  External  surface  smooth.  Section  shows 
wall  to  be  about  one-quarter  inch  thick.  Uterine  cavity  enlarged.    Con- 


THE  FEMALE  REPRODUCTIVE  ORGANS     305 

siderable  bloody  pus  in  uterine  cavity.  No  communication  can  be  found 
between  the  bladder  and  the  vaginal  wall  or  between  the  vaginal  wall 
and  the  rectum.  The  intestines  are  negative.  Mesenteric  glands  en- 
larged and  caseous. 

Abortion  and  miscarriage  are  occasionally  seen  in  the 
monkeys,  rodents  and  migulates.  The  underlying  reason 
for  this  can  seldom  be  determined.  A  few  cases  seem  to 
depend  upon  annoyance  or  abuse  by  cage-mates,  a  few  to 
immaturity  of  the  mother  and  some  others  seem  the 
result  of  bone  diseases  such  as  osteomalacia.  It  does  not 
occur  regularly  in  any  group  or  enclosure  and  appears  to 
have  no  comparative  pathological  value. 

Injuries  of  the  external  genitalia  of  the  nature  of, 
lacerated  wounds  are  occasionally  seen  in  ungulates  and 
carnivores,  but  never  in  monkeys.  Acute  vaginitis,  some- 
times localized  into  an  abscess  of  the  wall  has  been  known 
to  follow  these  traumata.  Chronic  changes  have  been 
met  but  once,  which  example  will  be  detailed  under 
another  heading. 

Inflammations. 

Endometritis  has  been  observed  sixteen  times,  as  fol- 
lows :  Carnivora  10,  Rodentia  3,  Ungulata  2,  Edentata  1. 
In  searching  for  causes  it  was  found  that  the  association 
with  a  recent  delivery  of  young,  an  abortion  or  the  reten- 
tion of  a  fetus,  was  responsible  five  times.  Association 
with  tumors  of  the  uterus  was  noted  four  times.  Injury 
preceded  the  condition  on  two  occasions,  while  one 
instance  seemed  to  be  hematogenic,  being  secondary  to  a 
septic  pneumonia.  On  four  occasions  the  actual  cause 
could  not  be  established  with  satisfaction.  Pathologically 
the  traumatic  and  parturient  cases  were  purulent  while 
tumors  seemed  to  produce  a  more  exfoliative  or  hyper- 
trophic inflammation. 

Inflammation  of  the  Fallopian  tube  has  been  observed 
but  five  times,  and  only  in  one  of  these  did  the  uterine 
wall  fail  to  participate  in  the  disease.     The   animals 


306   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

affected     were     three     carnivores,     one     rodent     and 
one  marsupial. 

No  particularly  important  association  of  this  sal- 
pingitis and  other  pathology  was  noted,  and  indeed  the 
only  noteworthy  lesion  of  the  organ  under  discussion  is 
now  to  be  cited. 

Nylghaie  (Boselaphus  tragocamelus) .  Chronic  vaginitis,  fibroma 
uteri,  chronic  tuberculous  salpingitis.  The  vulva  and  lower  half  of  the 
vagina  are  covered  by  a  thin  coating  of  yellowish  mucus.  Mucosa  is 
mottled  purple,  irregular,  in  some  places  smooth  and  flat,  and  in  others 
showing  clear  cysts  with  yellowish  fluid  contents.  The  upper  part  of 
the  vagina  shows  a  pedunculated  fibroid  extending  from  the  cervix. 
This  tumor  measures  32  x  20  mm.,  and  is  attached  to  the  posterior 
cervical  wall  by  a  broad  pedicle  and  is  of  smooth  surface.  The  right 
uterine  wall  shows  a  large  fibroid  which  twists  the  uterine  lumen  to  the 
left.  The  tumor  in  the  right  uterine  cornu  is  nodular,  measures  11  x  9 
X  7  cm.,  shows  many  dilated  veins  coursing  over  the  surface  and  one 
section  shows  a  soft  central  necrotic  area.  The  left  uterine  cornu 
shows  a  small  fibroid  at  the  lower  end.  There  is  a  chronic  endometritis 
present.  The  outer  half  of  the  right  tube  is  the  seat  of  firm  nodular 
swellings,  one  of  which  proves  to  be  a  purulent  salpingitis  (tubercu- 
lous), the  others  calcareocaseous  salpingitis.  Both  ovaries  are  fibroid 
and  cystic.  Microscopic  section  shows  some  exfoliation  of  the  epithe- 
lium of  the  vagina.  The  uterine  cornu  is  thickened  and  the  fibrous 
tissue  is  greatly  increased.  The  tube  is  the  seat  of  a  purulent  salpingitis 
with  an  occasional  broken  down  tubercle  present.  The  ovaries  show  a 
mild  degree  of  oophoritis   (Fig.  23). 

Obstructions  to  Continuity  of  Birth  Canal. 

Obstruction  to  the  lumen  of  the  genital  canal  was 
observed  in  three  mammals. 

An  American  Bison  {Bison  bison),  five  years  old  and  known  to  have 
been  in  captivity  three  years  died  of  a  chronic  gastroenteritis  and 
pulmonary  parasites  after  many  weeks  of  failing  health.  At  autopsy 
a  double  hydrosalpinx,  and  probably  unilateral  hydrometra,  Avith  low 
grade  cystic  oophoritis  were  found.  A  picture  is  rei^roduced.  The 
notes  are  not  perfectly  clear  as  to  the  anatomy  but  from  the  gross  speci- 
men in  preservative  it  would  seem  that  the  left  uterine  cornu  was 
involved  in  the  dilatation  (Fig.  21). 

Lion  cub  (Felis  leo)  had  congenital  cystic  uterus.  The  only  abnor- 
mality found  is  in  connection  with  the  internal  genitalia.  The  vagina 
is  50  mm.  long,  uterus  23  mm.  long,  horns  of  uterus  each  90  mm.  long. 


Fig.  2?.— PKDLNCL'LATED  FIBROMA  OF  CERVIX;  INTRAMURAL  FIBROMA  OF  UTERINE  BODY; 
FIBROMA  OF  LEFT  UTERINE  CORNU;  TUBERCULOUS  SALPINGITIS  AND  MURAL  METRITIS  RIGHT  SIDE. 
NYLGHAIK  (BOSELAPHUS  TRAGOCAMELUS). 


THE  FEMALE  REPRODUCTIVE  ORGANS     307 

The  diameter  of  the  uterus  measures  10  mm.  in  both  body  and  cornua, 
the  latter  being  distended  by  clear  watery  fluid.  They  are  symmetrically 
curved  downward.  They  show  no  adhesions  or  obliteration  of  the  os 
uteri  to  account  for  fluid  retention.  The  probe  is  readily  passed  from 
the  vagina  to  the  end  of  the  uterine  horns.  Vagina  is  patulous  through 
its  whole  course.     Ovary  and  tube  normal. 

An  Axis  Deer  {Cervus  axis)  showed  congenital  hydrometra.  This 
specimen  is  from  a  day  old  animal,  consists  of  the  genitalia  in  which 
the  entire  body  of  the  uterus  and  both  uterine  cornua  have  been  trans- 
formed into  a  thin  walled  translucent  cystic  cavity  containing  clear 
fluid.  The  body  of  the  uterus  measures  15  mm.  diameter,  each  cornu 
reaching  the  diameter  of  5  mm.  The  cornua  are  curled  not  unlike  ram's 
horns,  and  are  united  by  a  line  of  adhesions  above  the  body  in  the  mid- 
line. From  the  ends  of  the  convoluted  and  cystic  uterus  and  cornua 
spring  the  normal  tubes  each  with  its  ovary. 

Cysts. 

Cysts  have  not  been  noticed  in  the  lower  genital  canal, 
but  it  is  true  that  this  region  is  not  exhaustively  studied 
in  routine  autopsies ;  however,  no  large  cysts  have  occur- 
red there.  Cysts  of  the  ovary  have  been  limited  to  the 
so-called  cystic  disease  of  this  organ ;  one  parovarian  cyst 
has  been  found.  These  animals,  two  ungulates,  two  car- 
nivores and  a  rodent,  are  noted  as  being  young  adults, 
only  one  of  which  was  known  to  have  borne  young.  In 
only  one,  the  rodent,  was  there  evidence  of  chronic  peri- 
toneal disease.  The  notes  of  the  parovarian  cyst  case 
are  as  follows : 

An  Aoudad  {Ovis  tragelaphus)  showed  a  ruptured  parovarian  cyst 
with  normal  right  side  pregnancy.  The  fetus,  its  placenta  and  mem- 
branes show  no  change.  The  right  broad  ligament  and  ovary  are  nega- 
tive, the  latter  containing  a  red  and  gray  corpus  luteum.  The  left  ovary 
is  flbrotic  and  the  left  ligament  is  the  seat  of  a  large  hematoma,  which 
on  section  is  found  to  contain  thin  sheets  of  gi'ay  translucent  membrane, 
like  walls  of  a  cyst.  The  vessels  are  all  distended.  Arteries  have  stiff 
walls  and  are  empty.  Veins  have  well  formed  clot.  The  whole  uterus 
and  adnexa  were  slightly  turned  to  the  right,  but  the  twist  does  not 
seem  to  have  been  sufficient  to  cause  rupture  of  a  broad  ligament  vessel. 
Microscopic  section  of  the  broad  ligament  shows  a  parovarian  cyst  into 
which  there  has  been  hemorrhage.  The  cyst  is  separated  from  the  ovary 
proper  by  a  short  band  of  tissue  which  apparently  consists  of  thinned- 
out  ovarian  cortex. 


308   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Tumors. 

Tumors  of  this  tract  in  the  mammals  have  been  con- 
fined to  the  uterus  proper,  none  having  been  found  in  the 
vagina,  tubes  or  ovaries;  in  birds  one  oviductal  growth 
and  two  ovarian  tumors  are  recorded.  These  tumors  are 
reported  briefly  according  to  their  histological  structures, 
a  brief  summary  being  added  at  the  end. 

A  Black  Lemur  {Lemur  macaco)  showed  a  local  leiomyofibromatous 
nodule  on  the  lateral  aspect  of  the  uterine  body  near  the  cornu. 

A  nine-banded  Armadillo  (Tatu  novemcinctus)  gave  an  interesting 
specimen  which  can  be  described  as  follows:  The  uterus  is  enlarged 
so  that  it  measures  90  mm.  from  external  os  to  fundus.  Tubes  and 
ovaries  apparently  normal.  There  is  considerable  grumous  blood  in  the 
vagina  and  cervix ;  the  former  is  normal.  The  cervix  is  pale  and  opaque 
in  its  lower  half ;  upper  half  is  slightly  congested  and  mucosa  decidedly 
rugous.  The  utenis  itself  shows  an  attenuated  muscular  wall  with  a 
thickened  irregular  mucosa  which  is  the  seat  of  pseudomembranous  tabs 
of  a  dull  red  color,  while  the  mucous  membrane  itself  is  irregularly  red 
and  yellow;  also  some  grumous  blood  in  the  cavity.  The  size  of  the 
uterus  is  due  to  a  large  fibroma  attached  to  the  left  lateral  wall  near 
the  cornu.  The  mucous  membrane  of  this  is  irregularly  disturbed  in 
some  places,  the  tumor  being  partly  bare.  Here  and  there  the  mucous 
membrane  shows  the  same  degenerating  hypertrophic  character  as  seen 
on  uterine  wall.  The  tumor  is  attached  to  the  wall  by  a  narrow  pedun- 
cle (Fig.  25). 

A  nylghaie  with  multiple  fibromata  has  already  been  described. 

A  Jaguar  {Felis  onea)  presented  in  the  middle  of  the  right  uterine 
cornu  a  cystic  resilient  tumor  3x4  cm.,  which  proved  to  be  a  fibro- 
adenoma. 

A  lioness  (Felis  leo)  presented  a  penetrating  malignant  adenoma  of 
the  uterine  cervix  upon  which  an  active  endometritis  was  implanted. 
The  tumor  penetrated  the  uterine  wall,  which  gave  way,  an  acute  fatal 
peritonitis  resulting.    Metastases  had  occurred  to  the  lung. 

A  Wild  Boar  {Sus  scrofa)  had  a  generalized  ulcerating  carcinoma 
of  the  uterine  body. 

An  instance  of  chorion-epithelioma  in  a  Canada  porcupine  is 
worthy  of  separate  description. 

Canada  Porcupine  (Erethizon  dorsatus  dorsatus).  Acute  suppura- 
tive catarrhal  endometritis,  hemorrhage  in  myometrium,  chorion-epithe- 
lioma. In  anterior  abdominal  wall  a  short  quill  was  found  imbedded. 
A  very  small  quill  Avas  imbedded  in  the  retrocervical  muscles.  A  quill 
about  2  cm.  long  lies  free  in  the  peritoneal  cavity  attached  by  recent 
plastic  adhesions  to  peritoneum  over  left  pubic  ramus.  A  fourth  quill 
was  adherent  by  recent  fibrous  lesions  to  anterior  wall  of  cecum.  Peri- 
toneum contains  a  moderate  excess  of  thin  watery  fluid.     Liver  and 


p,r.  '>6— FIBROMYOMA  OF  UTERUS.  CORNUA  AND  TUBES.  INDIAN  ELEPHANT  (ELEPHAS 
INDICUS)."  ■  WHOLE  MASS  AS  MOUNTED  ON  BOARD.  IT  MEASURED  WHEN  FRESH  ABOUT  SIX 
I-EET  ACRObS_^^^^^  SECTION  OF  CORNU  OF  Fir..  26  AT  HIGHER  POWER.  A  PIECE  CUT  OFF 
WHERE  THE  PALE    AREA  SHOWS  ON  THE    RIGHT  CORNU  OF  THE  OTHER   FKJURE. 


THE  FEMALE  REPRODUCTIVE  ORGANS  309 

spleen  are  negative.  Kidney  is  large,  greenish  yellow,  firm,  smooth  and 
glistening.  The  right  uterine  cornu  is  subinvoluted  (the  animal  was 
delivered  of  a  fetus  before  arrival  at  the  Garden,  which  was  sixteen 
days  before  death).  Its  walls  are  thick  and  distinctly  congested,  the 
congestion  being  of  inflammatory  type.  Microscopic  section  of  uterus 
shows  a  regular,  not  ulcerated  serosa.  Subjacent  fibrous  tissue  is  loose 
and  contains  a  granular  precipitate  together  with  a  few  red  blood  cells. 
Muscular  bundles  under  this  are  widely  separated  evidently  partly  by 
trauma,  but  certainly  also  by  edematous  interstitial  tissue  in  which 
fibrillcB  are  widely  separated  and  between  which  free  red  blood  cells 
and  plasma  cells  are  seen.  Capillaries  ramifying  through  muscular 
bundles  are  greatly  distended  and  congested.  Numerous  large  arteries 
are  present  in  addition.  Some  of  these  contain  pink  granular  material 
within  their  walls  together  with  diffuse  collection  of  red  blood  cells. 
Fibroblasts  extend  into  this  necrotic  mass  from  other  sections  of  the 
walls.  Lumen  of  such  arteries  is  diminished  and  in  places  quite  obliter- 
ated by  recent  organization  tissue.  At  many  places  in  muscularis  are 
large  cells  of  iiTegular  rounded  form  and  some  cytoplasms  incline  to- 
ward the  basic  tint  with  one  or  several  large  hyperchromatie  nuclei. 
They  are  especially  likely  to  occur  close  to  a  capillary.  There  is  an  es- 
pecially large  accumulation  of  these  cells  at  that  point  of  section 
farthest  from  fundus.  Here  these  cells  occur  in  chord-like  masses 
which  infiltrate  the  muscularis  both  internally  and  externally.  This 
particular  mass  lies  in  the  muscularis  internal  to  great  arteries  and 
well  removed  from  mucosa.  In  this  mass  are  giant  cells  with  multiple 
nuclei  scattered  through  the  whole  cytoplasm  together  with  smaller 
cells  Avith  exceptionally  large  hyperchromatie  nuclei.  Subepithelial 
tissue  is  especially  congested  and  contains  numerous  fibroblasts  together 
with  a  few  well  formed  glandular  acini.  Lining  epithelium  is  discon- 
tinous,  of  simple  tall  columnar  type,  in  places  becoming  flattened  or 
even  lost.  Lumen  of  organ  is  practically  completely  occupied  by  pus. 
The  Indian  Elephant  "  Empress  "  (Elephas  indicus)  showed  calci- 
fied fibroids  of  the  fimbriae;  gross  and  microscopic  notes  and  a  photo- 
graph are  given.  The  uterus  is  bicornate  in  type.  In  its  body  there  are 
numerous  fibrous  nodules  1  to  4  cm.  diameter.  They  can  be  traced  from 
the  cervix  to  the  ends  of  both  horns  and  tubes.  At  the  end  of  each 
tube  there  is  a  gi*eat  mass  of  calcified  partly  conglomerate  tumors  some 
of  which  are  partly,  others  quite,  pedunculated.  One  specimen  measur- 
ing 2xlxli  cm.  has  a  peduncle  15  cm.  long.  The  mass  on  the  right  side 
weighs  3,926  grams,  that  on  the  left  side  about  the  same.  Section  of 
tumor  from  uterine  cornu  shows  the  classical  appearance  of  a  leiomyoma 
with  usual  whorling  and  interlacing  bundles  of  involuntary  muscle 
fibres.  Degenerative  and  vascular  changes  not  seen  nor  is  there  any 
notable  addition  of  fibrous  tissue.  This  latter  tissue  is  shown  only  in 
small  amounts  at  one  end  of  section. 

The  preceding  data  record  the  discovery  of  three 
fibromata  of  the  uterus  and  one  of  the  fimbriae ;  one  fibro- 


310   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

adenomata  of  the  cystic  type,  one  malignant  adenoma, 
one  adenocarcinoma,  and  one  chorion-epithelioma.  They 
were  found  in  Lemures  1,  Carnivora  2,  Ungulata  2,  Pro- 
boscidea  1,  Rodentia  1,  and  Edentata  1.  Metastases 
occurred  but  once,  to  the  lung.  No  secondary  tumors  were 
found  in  the  uterus. 

While  upon  the  subject  of  neoplasms  of  the  female 
genitalia,  it  may  be  well  to  describe  the  three  instances 
found  in  the  avian  organs.  A  very  definite  case  of  adeno- 
carcinoma occurred  in  the  oviduct  of  an  Undulated  Grass 
Parrakeet  (Melopsittacus  undulatus),  a  variety  of  bird 
very  susceptible  to  neoplasms.  The  notes  in  an  abbre- 
viated form  are  given : 

Immediately  under  the  ovary  is  an  irregular  mass  measuring  2  cm. 
long,  1  cm.  wide,  1  cm.  deep.  The  lower  part  of  the  tumor  thus  comes 
to  press  against  the  cloaca.  It  is  adherent  anteriorly  to  the  peritoneum. 
It  apparently  consists  of  two  parts,  an  upper  rounded  larger,  and 
a  lower  spherical  smaller.  Both  parts  are  well  encapsulated 
and  separated  from  each  other  by  a  well  defined  constriction. 
The  upper  part  has  a  pale  opalescent  appearance.  It  cut  easily  with 
moderate  resistance.  The  lower  portion  externally  has  an  egg-yellow 
color  streaked  with  red.  Upon  section  it  has  the  same  general  appear- 
ance but  contains  in  addition  numerous  small,  irregular,  yellow  areas 
which  mask  the  general  opalescent  appearance.  The  centre  of  this  node 
contains  an  empty  space  (cyst)  1x2  mm.  Microscopic  section  consists 
of  an  oval  or  elliptical  mass  showing  over  one  convexity  a  depression 
simulating  a  constriction.  A  thin  fibrous  capsule  extends  over  most  of 
the  section  which  is  extra  thick  at  the  point  of  constriction.  Constriction 
roughly  divides  the  section  into  two.  The  upper  portion  consists  of 
one  or  two  coarse  septa  of  fibrous  tissue.  From  these  central  areas  a 
delicate  connective,  tissue  framework  extends  peripherally.  In  this 
framework  are  great  numbers  of  irregular  gland  spaces.  These  gland 
spaces  are  so  closely  placed  in  most  cases  that  room  is  afforded  for  but 
one  nucleus  of  the  bundle.  The  gland  spaces  vary  in  size,  some  large, 
some  small,  and  show  grotesque  shapes.  The  larger  gland  spaces  here 
contain  granular  debris  and  pyknotic  nuclei.  Compound  granule  cells 
suggesting  colostrum  corpuscles  may  be  seen  in  this  debris.  The  epi- 
thelium of  the  gland  spaces  consists  of  a  single  layer  of  columnar 
epithelium  of  low  cuboidal  type.  In  places  it  is  heaped  up  so  as  to 
present  several  layers.  In  places  too  it  is  not  applied  in  a  regular 
manner  to  the  basement  membrane  but  breaks  through  and  then  the 
cells   extend   in   most   disorderly   fashion   into   the   lymphatics   of   the 


Fk;.   28. — PAPILLARY  ADENOMA  OF  OVARY.     WILD  TIRKKY 
(MELLEAGRIS   GALLOPAVO). 


THE  FEMALE  REPRODUCTIVE  ORGANS  311 

stroma.  At  these  points  the  nuclei  are  hyperchromatie.  The  lower 
portion  follows  closely  the  description  given  above  save  that  the  glandu- 
lar spaces  are  much  larger.  They  contain  pink  glandular  material  Avith 
admixture  of  compound  granule  cells.  At  the  convexity  of  the  tumor 
the  acini  are  especially  large.  Here  they  contain  a  pink  granular  mate- 
rial which  stains  more  intensely  than  the  other  granular  contents  and, 
too,  inside  of  this  intense  pink  material  are  sharply  circumscribed 
areas  of  yet  more  intensely  pink  staining  material.  This  latter  sub- 
stance has  a  streaming  appearance  under  the  high  power.  This  stream- 
ing appearance  is  due  to  elongated  areas  of  less  dense  material  which 
are  placed  with  their  long  axes  parallel.  This  lower  portion  shows, 
furthermore,  even  with  the  naked  eye,  two  large  cysts  which  are  lined 
by  epithelium  and  contain  a  very  small  number  of  compound  granule 
cells.  The  capsule  at  the  lower  polq  is  worthy  of  note  from  the  extreme 
dilatation  of  its  capillaries. 

A  papillary  adenoma  was  found  in  a  wild  turkey 
(Fig.  26)  and  a  mixed  cell  sarcoma  in  a  King  parrakeet 
{Apromictus  cyanopygius).  None  of  these  tumors  sent 
out  metastases. 

Interest  in  the  avian  reproductive  tract  from  a  patho- 
logical standpoint  centres  around  the  tumors  as  already 
given,  and  abnormalities  in  egg-bearing.  Among  our 
specimens  there  have  been  many  cases  of  soft  shelled  eggs 
apparently  blocked  in  the  oviduct,  of  ''egg-binding"  and 
of  the  inspissated-egg-remains  in  the  abdominal  cavity. 
These  conditions  are  well  known  to  veterinarians  and  are 
explained  on  the  basis  of  improper  food,  immaturity  of 
the  bearing  fowl,  injury  and  inflammations  of  the  cloaca 
and  oviduct.  I  made  an  attempt  to  associate  these  con- 
ditions with  infectious  disease  incidence  and  with  the 
normal  egg  size.  The  results  are  not  harmonious.  No 
relation  existed  between  general  or  local  infection  and 
any  of  these  conditions.  Gallinaceous  birds  with  their 
large  eggs  show  the  highest  percentage  (2.3  per  cent.), 
but  Anseres  witli  a  somewhat  larger  relative  egg  size 
show  1.6  per  cent.  Passeres,  with  eggs  of  very  variable 
size  but  relatively  large  pelves,  have  an  incidence  of  .6  per 
cent.  Struthiones'  eggs  are  relatively  small;  their  inci- 
dence is  1.5  per  cent.  \ 


312   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Salpingitis  occurred  in  seven  cases,  but  there  have 
been  no  evidences  of  an  acute  infectious  disease  such  as  is 
responsible  for  gleet. 

The  Mammaby  Gland. 

This  structure  has  been  the  seat  of  three  inflammatory 
conditions  and  four  tumors.  The  former  occurred  twice 
in  nursing  animals,  one  of  which  seemed  to  be  suffering 
with  distemper,  another  from  puerperal  sepsis,  while  the 
third  case  was  probably  traumatic.  The  animals  were  in 
order,  a  raccoon,  a  coati  and  a  skunk.  Four  cancers  of  the 
breast  have  been  observed,  all  richly  cellular  or  glandular 
in  type;  none  of  the  scirrhous  variety  has  been 
encountered.  Two  of  them  were  ulcerating  and  one  was 
about  to  become  so.  One  gave  extension  to  the  axilla  and 
lung,  one  had  penetrated  the  abdominal  wall  and  growm 
around  the  kidney ;  the  remainder  had  not  yet  extended. 
The  animals  were  Black  Bear  {Ursus  americanus) 
(thoracic  mamma  with  extension),  Common  Opossum 
(Didelphys  virginiana)  (two  sections  of  breast  in  pouch), 
two  White-footed  Mice  {Peromyscus  leucopus)  (posterior 
abdominal  right  gland  and  whole  side).  These  animals 
were  all  adult  but  not  old. 


29. — ADENOCARCINOMA   OF  MAMMARY  GLAND    (TWO  SECTIONS   WITHIN   POUCH). 
COMMON   OPOSSUM    (DIDELPHYS  VIRGINIANA). 


SECTION  X 
THE  MALE  GENITALIA 

Affections  of  the  penis  are  limited  to  phimosis  and 
occasional  inflammations  on  a  traumatic  basis.  The  lat- 
ter is  best  seen  in  carnivorous  animals  like  raccoons  and 
coatis,  but  has  little  pathological  interest.  Two  cases  of 
phimosis  have  been  seen  at  the  autopsy  table  and  one  in 
an  animal  (hyena)  still  living.  The  last  was  operated 
upon  some  years  ago  and  has  given  no  trouble  since.  The 
prepuce  of  this  beast  is  seldom  retracted,  but  no  swelling 
or  retention  of  urine  has  occurred.  A  Eed  River  Hog 
{Potamochoerus  porcus)  dying  of  a  variety  of  lesions, 
was  found  to  have  contracted  preputial  opening,  the 
edges  of  which  were  tight  and  adherent  to  the  glans  penis 
at  various  spots.  The  prepuce  had  been  dilated  with  urine 
to  a  large  size  from  which  collection  the  fluid  could  be 
pressed  dropwise  only  by  considerable  pressure.  Opening 
the  sac  revealed  forty  to  fifty  gray  white  sand  granules 
about  the  size  of  millet  seeds.  A  gray  fox  {Canis  cinereo 
argenteus)  had  a  mild  grade  of  phimosis;  in  this  case 
probably  traumatism  had  some  etiological  relation,  for  it 
is  the  one  referred  to  before  in  which  a  terminal  urethral 
stricture  was  followed  by  rupture  of  the  bladder. 

The  testes  have  been  peculiarly  free  of  disease,  only  a 
small  number  of  lesions  having  been  found.  A  raccoon  had 
an  acute  inflammation,  traumatic  in  origin,  and  a  few 
passerine  birds  were  noted  as  shomng  involvement  of  this 
organ  in  the  presence  of  some  general  infectious  diseases. 
( Two  tumors  were  found  in  birds,  none  in  mammals.  The 
avian  cases  both  occurred  in  Red-shouldered  Parrakeets 
{PalcBornis  eupatrius)  and  were  round  cell  sarcomata, 
without  metastases. 

Acute  inflammations  of  the  prostate  and  Co^vper's 
glands  occurred  only  once  as  secondary  to  pelvic  infec- 

21  313 


314   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

tion.  Chronic  change  was  observed  in  these  bodies  on  two 
occasions,  and  prostatic  hyperplasia,  commonly  called 
hypertrophy,  was  seen  thrice.  The  mammalian  prostatic 
area  is  knowTi  to  become  overfilled  with  secretion  and  to 
be  affected  by  inflammation  when  it  is  not  discharged. 
This  occurred  in  a  wood  rat  {Neotoma  pennsylvanica) 
and  a  wild  boar  (Sus  scrofa),  the  former  having  the 
condition  so  marked  that  prostate  and  seminal  vesicles 
were  swollen  backward  into  the  pelvis  like  a  tumor.  His- 
tologically one  finds  in  these  conditions  an  engorgement 
of  the  glandular  and  ductal  spaces  with  a  mucoid  sub- 
stance and  a  mild  round  and  plasma  cell  infiltrate  in  the 
connective  tissue;  there  are  no  marked  evidences  of 
active  inflammation.  The  testes  were  not  abnormal  in 
these  cases. 

The  three  cases  of  ''hypertrophy"  of  the  prostate  are 
worthy  of  separate  record. 

Indian  Paradoxure  {Paradoxurus  niger)  the  prostate  is  greatly 
enlarged,  of  pale  orange  color,  soft  and  does  not  exude  pus  on  section 
or  pressure. 

Common  Opossum  {Didelphys  virginiana)  The  prostate  is 
enlarged,  soft,  gray  yellow.  Urethra  contains  a  little  gray  mucus  but 
seems  patulous.     Seminal  vesicles  negative. 

Rhesus  Macaque  (Macacus  rhesus).  Glandular  hyperplasia  of 
prostate.  The  bladder  is  collapsed.  The  opening  of  the  urethra  is 
occupied  by  a  firm,  friable  yellow-white  cast  of  matter  apparently  in- 
spissated semen.  Urethral  mucous  membrane  nonnal.  Prostate  is  7  x 
4.5  X  3  cm.  firm,  resilient,  dull  purple-gray,  capsule  negative.  Section 
shows  normal  lobulations  containing  apparently  normal  secretion.  Vasa 
negative.  Testes  negative.  Microscopic  section  of  prostate  shows  acini 
of  various  sizes  lined  by  a  single  layer  of  tall  vacuolated  cells  with 
nucleus  at  the  bottom.  Cells  probably  nowhere  reduplicated.  Acini 
nowhere  grossly  atypical  but  everywhere  hyperplastic  and  dilated  irregu- 
larly. Mass  is  adenomatous  in  general  increase  but  no  part  is  truly 
neoplastic.  Interstitial  tissue  rather  less  than  normal  proportion- 
ately.   No  "  amyloid  "  bodies. 

Still  another  case  of  accessory  sex  gland  enlargement 
was  found  in  a  Ring-tailed  Lemur  {Lemur  catta),  this 
time,  however,  with  more  suggestion  of  a  neoplastic 
change.     The  prostate  of  the  lemur  is  normally  large. 


IHK  PROSTATE  CJI 


THE  MALE  GENITALIA  315 

pale  gray-pink  and  rather  firm.  Histologically  it  is  about 
equally  glandular  and  fibrous.  In  this  case  the  cellular 
activity  is  undoubted,  and  one  must  consider  it  adenom- 
atous. The  cause  of  death  was  enteritis,  being  perhaps 
more  serious  in  the  presence  of  the  urethral  obstruction. 

The  seminal  vesicles  are  distended  to  7  x  2  cm.  with  a  thick  boiled- 
starch-like  material.  Wall  and  mucosa  are  negative.  Prostate  large, 
tense  and  injected.  Its  cut  surface  is  pale  purple,  homogeneous; 
character  of  fluid  is  normal  although  excessive.  Urethra  is  occupied 
by  a  cast  of  rather  tenacious  starch-like  matter  which  begins  at  neck  of 
bladder  and  runs  almost  to  meatus.  Openings  of  excretory  ducts  are 
prominent.  Mucosa  and  submucosa  of  urethra  are  deep  purple  and  the 
former  seems  to  be  slightly  opaque  as  if  covered  with  desquamated 
epithelium.  Testes  and  epididymes  seem  normal.  Vasa  deferentia  are 
slightly  distended  with  excess  of  normally  turbid  fluid.  Microscopical 
section  shows  hyperplastic  epithelial  condition  with  accumulation  of 
droplets  of  hyaline  matter  but  there  is  no  amyloid  deposit.  In  places 
it  is  possible  to  see  a  hyperplastic  and  loosened  epithelium  with  nuclei 
becoming  vacuolated,  and  the  whole  being  cast  off.  Less  granular  free 
globules  suggest  that  this  is  the  method  of  origin  of  the  hyaline  globules 
free  in  the  acini.  The  picture  is  one  of  papillary  adenoma.  In  some 
places  there  is  surely  reduplication  of  the  lining  cells.  Connective 
tissue  is  deeply  staining,  compact  and  with  adult  nuclei.  Growth  is  not 
very  vascular.    There  are  no  corpora  amylacea. 

A  case  of  tuberculous  prostatitis  and  seminal  vesic- 
ulitis was  seen  in  a  Japanese  Macaque  {Macaciis  fusca- 
tus).  Judging  b}^  the  advanced  stage  of  these  lesions  and 
their  more  recent  character  in  other  viscera,  the  disease 
was  suspected  as  pelvic  in  origin,  possibly  due  to  infection 
by  a  thermometer.  Whether  or  not  such  be  the  case  cannot 
be  established,  but  at  all  events,  separate  thermometers 
kept  in  carbolated  vaseline  were  employed  after  this 
death.  The  females  caged  with  this  animal  did  not  de- 
velop tuberculosis  of  the  pelvic  organs. 


SECTION  XI 
THE  DUCTLESS  GLANDS 

The  Thyroid  Bodies. 

The  ductless  glands  occupying  the  anterior  cervical 
regions,  knowoi  as  the  thyroids  and  parathyroids  are 
structures  to  be  found  in  some  form  in  all  vertebrates, 
but  increase  in  distinctness  of  outline  and  construction 
upward  in  the  zoological  scale.  In  the  bird  they  present 
themselves  as  discrete  rounded  bodies  lying  well  to  the 
side  of  the  midline  resting  usually  upon  the  carotid  artery 
or  jugular  vein  or  both.  As  one  removes  the  skin  reddish 
brown  globular  masses  will  be  exposed  to  view,  sometimes 
showing  an  irregular  lobular  outline,  a  variation  due  to 
separate  but  attached  masses  of  parathyroid  glandules ; 
the  latter  may  be  yellowish  or  even  white.  For  the  most 
part,  however,  distinct  thyroid  and  parathyroid  bodies 
are  separated  with  difficulty,  and  one  must  discover  the 
latter  by  microscopical  section.)  In  the  mammal  these 
organs  are  by  no  means  so  readily  found  when  removing 
the  cervical  integument,  for  they  are  usually  buried 
beside  the  trachea  and  covered  by  sternohyoid  and 
sternothyroid  muscles.  Their  position,  relative  to  the 
larynx  and  upper  end  of  the  sternum,  varies  considerably 
but  this  seems  to  have  little  importance  in  the  enlarge- 
ments to  which  the  gland  is  liable.  It  is,  however,  lower, 
that  is  more  posterior,  than  in  the  human  being,  rarely 
rising  as  high  as  the  lateral  thyroid  cartilages.  The 
principal  lobes  are  elliptical  or  roughly  triangular  masses 
with  their  long  axis  corresponding  to  that  of  the 
animal 's  body  and  apposed  mesially  to  the  trachea.  AVhen 
enlargement  occurs  it  develops  in  the  anterior  or  ventral 
direction,  pushing  through  the  cleft  between  the  muscles 
and  the  trachea  to  present  under  the  cervical  skin.    In 

316 


THE  DUCTLESS  GLANDS  317 

quadrupeds  it  may  become  dependent  and  the  swelling  is 
nearer  the  sternum  than  is  the  case  in  man.  The  isthmus 
is  a  very  variable  structure,  and  its  presence  or  absence 
cannot  be  said  to  be  a  constant  character  in  any  order,  or 
indeed  in  any  family.  I  have  seen  in  old  animals  a  fibrous 
band  extending  over  the  face  of  the  trachea  connecting 
the  capsule  of  the  lateral  lobes,  which  might  have  been 
an  isthmus  at  one  time.  From  these  few  observations  the 
idea  of  atrophic  fibrosis  occurred  to  me.  Such  may  be 
the  reason  for  the  absence  of  this  transverse  link  in  some 
adult  specimens. 

The  amount  of  thyroid  tissue  possessed  by  an  animal 
might  be  judged  by  measurement  or  weight.  The  former 
is  misleading  since  the  density  might  vary,  as  it  certainly 
does  in  the  two  classes  and  between  certain  orders  in 
mammals.  Actual  weights  would  afford  little  comparison, 
whereas  the  weight  in  terms  of  total  body  weight  may 
supply  a  guide  to  the  amount  of  gland  normal  to  an 
animal.  There  are  given  in  Table  15  the  grams-per-kilo- 
gram-body- weights  of  the  thyroid  bodies  (thyroid  and 
parathyroids  both  sides  combined)  of  twenty  animals 
whose  gland  seemed  entirely  normal  at  autopsy.  They 
are  all  adult  specimens,  free  of  cretinoid  characters  and 
of  bone  or  heart  diseases,  conditions  which  might  reflect 
abnormalities  to  these  glands.  The  list  is  too  small  to 
warrant  any  conclusions,  but  in  one  respect  confirms 
Murray's  (1)  observation  and  certain  experimental  work, 
notably  of  Vincent  and  Jolly  (2)  and  Carlson,  Rooks  and 
McKie  (3).  The  carnivores  have  more  thyroid  than 
ungulates  (averages  .55  gm.  vs.  .18  gm.),  but  the  mar- 
supials on  our  list  have  nearly  as  high  an  average  as  the 
former,  namely  .44  gm.  To  these  figures  might  be  added 
others  which  I  have  worked  out  from  the  list  given  by 
Murray ;  it  is  only  possible  to  compute  the  gram-per- 

(1)  Proc.  L.  Z.  8oc.,   1919,  p.  16. 

(2)  Journ.  Phys.,  Vol.  34,  295. 

(3)  Am.  Jour.  Phys.,   Vol.  30,  129. 


318   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Table  15 

Showing  Weights  of  Animals,  of  Their  Thyroid  Bodies  and  the  Relation  of  These 
Weights  Per  Kilogram. 


Grams  of  Body 


Grams  of 
Thyroid 


Grams  of 
Thyroid  per 
Kilo  of  Body 


Primates : 

tei  Woolly  Monkey 

Lagothrix  lagotricha 
Carnivora: 
Wild  Cat 

Felis  ruffus 
Silver  Fox 

Cams  chama 
Kamchatkan  Bear 

Ursus  beringiana 
Lynx 

Felis  canadensis 
Jaguar 

Felis  onca 
Rodentia: 
Ground  Squirrel 

Xerus  capensis 
Ungulata: 

Zebra 

Equus  burchelli 

Giraffe 

IKfeGiraffa  camelopardalis 
Barasingha  Deer 

Cervus  duvanceli 
Kashmir  Deer 

Cervus  cashmirianus 
Urial 

Ovis  vignei 
Marsupialia: 

Kangaroo 

Macropus  robustus  (?) 
Tasmanian  Devil 

Sarcophilus  ursinus 
Wombat 

Phascolomys  mitchelli 
Wallaby 

Genus  and  species  (?) 
Edentata: 

Anteater 

Myrmecophaga  tetradactyla 
Accipitres : 
Wedgetailed  Eagle 

Aquila  audax 
Struthiones: 

Rhea 

Rhea  americana 

Emu 

Dromseus  novae-hollandise 


2.370 


8,180 

2. 

3,325 

4. 

31,800 

22. 

9,500 

1 

29,500 

16 

550 

340,000 

384,000 

56,800 

56,800 

22,700 

18,000 
3,120 

26,000 
3,360 

3,300 

2,300 

18,000 
36,300 


45. 

8. 
20. 

2.5 

10. 
4. 

7. 
3. 

2. 

1. 

5. 
23. 


.84 

.24 
1.2 
.63 
.1 
.54 

3.6 

.19 
.12 
.14 
.35 
.11 

.55 
.13 
.27 
.9 


kilogram  value  for  a  few  of  his  examples  since  the  body 
weights  are  not  given  in  all.  Using  2.2  pounds  as  equal  to 


THE  DUCTLESS  GLANDS  319 

one  kilogram,  the  seal  of  432  kilos  had  .03  gm.  of  thyroid, 
lion  had  .18  gm.,  leopard  had  .46  gm.,  a  serval  .36  gm., 
a  skunk  2.35  gm.,  (was  this  normal?)  while  a  single 
herbivore  with  the  weight  given  was  a  porcupine  having 
.25  gm.  per  kilo.  The  average  of  Murray's  carnivores 
is  therefore  .67  gm.  per  kilo  of  body  weight,  whereas  our 
figure  is  .55  gm.  That  the  incidence  of  thyroid  abnor- 
malities stands  in  direct  relation  to  carnivorous  character 
has  been  recognized  before,  and  is  abundantly  borne  out 
by  our  statistics,  as  will  appear  at  a  later  place. 

Anatomy. 

The  minute  anatomy  of  the  normal  thyroid  is  fully 
given  in  text-books,  and  is  doubtless  pictured  in  the  minds 
of  all  but  students  of  the  subject  as  a  fixed  and  definite 
affair.  Such  is  not  the  case.  There  is  certainly  a  variation 
in  gross  size  under  conditions  of  seasonal  and  sexual 
activity,  and  it  would  seem  acceptably  demonstrated  that 
changes  in  diet,  especially  where  meat  is  concerned,  are 
associated  with  swellings  or  shrinkage  of  the  glands. 
These  gross  changes  must  be  due  to  alterations  in  his- 
tology. In  youth  also  the  bulk  is  larger,  a  condition  due 
to  cellular  activity,  while  as  middle  age  advances  the 
thyroid  becomes  smaller  and  more  balanced  in  its  colloid 
and  cellular  proportions.  I  need  not  detail  the  ultimate 
constituents  of  the  gland,  but  it  is  well  to  emphasize  a  few 
points  which  must  be  taken  into  consideration  in  micro- 
scopical diagnosis.  In  the  first  place,  the  lobules  or  acini 
are  not  all  of  the  same  size  in  normal  organs.  This  is 
especially  true  in  the  normal  adult  gland  but  may  be  so  in 
youth.  The  cells  which  form  the  inner  lining  of  the  acinus 
are  low  cuboidal  in  shape  but  the  elements  which  lie  under 
them  are  oval  and  do  not  change  with  hyperplasia  of  the 
former.  The  colloid  which  fills  the  glandular  spaces  is 
very  susceptible  to  mechanical  and  chemical  agencies.  In 
material  preserved  in  alcohol  it  may  be  shrunken  away 
from  the  cells   or  heavily  vacuolated,  while  in  tissue 


320  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

preserved  a  long  while  in  any  liquid  it  may  be  found  con- 
tracted or  broken.  Large  vacuolated  or  vesicular  cells 
are  characteristic  of  the  human  parathyroid ;  this  type  is 
by  no  means  so  prominent  in  the  lower  animals,  and  in  the 
few  examples  studied  closely  the  arrangement  is  less 
definitely  glandular  than  in  man. 

The  avian  thyroid  is  distinct  from  the  mammalian  in 
the  delicacy  of  its  fibrous  framework  and  the  flatter 
character  of  its  epithelia.  Acini  are  usually  of  more 
uniform  size.  Capsular  vessels  are  prominent  but 
internal  vascularity  is  less  in  birds  than  in  mammals  and 
their  goitres  are  not  solid. 

Physiology. 

The  physiological  value  of  the  thyroid-parathyroid 
complex  has  been  the  subject  of  extensive  study  and 
voluminous  literature  without  exhaustion  of  the  possi- 
bilities, but  with  the  result  that  we  are  possessed  of 
knowledge  explaining  certain  phases  of  abnormality,  even 
if  the  normal  functions  be  not  unexceptionally  demon- 
strated. The  accepted  alterations  of  functions  are 
hypothyroidism — inadequate  physiology,  and  hyperthy- 
roidism— excessive  activity.  Absence  or  atrophy  of  the 
thyroid  bodies  is  usual  in  hypothyroidism,  w^hile  enlarge- 
ments, collectively  called  goitre,  commonly  accompany 
excessive  function.  Exemplifying  the  former,  cretinism 
is  the  result  of  failure  of  normal  function  and  develop- 
ment during  fetal  life  while  myxedema  is  the  expression 
of  the  disappearance  of  thyroid  secretion  after  it  has  once 
been  operative;  the  latter  may  occur  in  infancy  after 
nursing  has  ceased,  or  at  any  time  that  the  thyroid  may 
atrophy,  during  some  of  the  forms  of  goitre  for  example. 
Hyperthyroidism  may  express  itself,  with  or  without 
visible  enlargement  of  the  thyroid  body,  in  nervousness, 
gastrointestinal  disturbances,  tachycardia,  loss  of  weight 
and  exophthalmos.  The  first  group,  which  might  be  called 
athyroidism,  is  often  associated  with  alterations  in  the 


THE  DUCTLESS  GLANDS  321 

bony  skeleton  in  the  form  of  chondrodystrophy  or  of 
rickets,  while  some  degree  of  osseous  change  is  observed 
with  myxedema.  Goitre,  be  it  simple  or  exophthalmic,  may 
be  foUow^ed  by  cardiac  enlargement  or  by  myocarditis. 

The  character  of  the  thyroid  secretion  is  not  kno^vn 
except  that  it  is  influenced  by  the  availability  of  iodine  in 
the  diet,  but  there  are  some  other  as  yet  unexplained 
features.  The  normal  thyroid  fed  to  cretins  or  persons 
suffering  with  myxedema,  has  the  power  to  improve  the 
condition  apparently  by  supplying  iodine  and  the  other 
essential  elements.  Iodine  is  an  important  constituent 
of  the  gland,  being  present  in  combination  with  protein.! 
Its  quality  varies  indirectly  with  the  amount  of  colloid 
and  of  hyperplasia  (Marine).  The  administration  of  this 
element  is  beneficial  in  colloid  goitre  but  is  harmful  in  the 
toxic  variety.  Even  though  the  administration  of  thyroid 
extract  may  relieve  athyroidism,  this  procedure  in  normal 
animals  fails  to  produce  typical  pictures  of  hyperthyroid- 
ism. Carnivora  fed  thyroid  gland  do  not  show  toxic 
symptoms  until  excessive  amounts  are  given,  whereas 
herbivorous  varieties  are  much  more  sensitive  to  this 
feeding. (4)  Tachycardia,  nervousness  and  exophthalmos 
are  not  produced  by  these  experiments,  an  interesting 
observation  since  these  signs  are  not  recorded  in  wild 
animals,  and  only  vaguely  reported  by  veterinarians. 
Man  is  apparently  very  sensitive  to  thyroid  dysfunction. 

On  the  other  hand,  meat-eating  animals  are  more 
sensitive  to  excision  of  the  thyroid  body  than  are  grain 
eaters. (5)  Chemical  studies  have  shown  that  the  thyroid 
is  concerned  in  basal  metabolism  since  this  is  increased 
in  hyperthyroidism  and  decreased  in  myxedema;  nitro- 
gen output  is  much  elevated. 

Thyroid  physiology  stands  probably  in  some  relation 
to  the  cardiac  mechanism  since  in  simple  colloid  or  simple 
hyperplastic  goitre  if  of  long  duration,  cardiac  enlarge- 

(4)  Carlson,  Rooks  and  McKie,  Loe.  cit. 

(5)  Vincent  and  Jolly,  Loc.  cit. 


322   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

ment  and  disease  may  be  greater  than  the  excess  work 
occasioned  by  the  mere  physical  bulk  of  the  enlarged 
gland  in  the  neck  would  seem  to  warrant.  A  detoxicating 
function  has  been  ascribed  to  the  gland,  but  Murray  dis- 
credits this  on  the  ground  that  congestion  occurs  in 
infectious  disease  of  warm  blooded  animals  but  not  in 
snakes.  He  thinks  the  gland  more  likely  stands  in  some 
connection  with  the  thermoregulatory  mechanism. 

It  is  evident  from  the  foregoing  that  the  thyroid  is 
closely  related  to  protein  metabolism,  and  that  this  is  in 
some  way  connected  with  the  ability  the  body  possesses  to 
use  iodine  in  the  food  if  it  can  get  it.  Feeding  of  meat  to 
fish  was  found  by  Marine  to  increase  the  size  of  the 
thyroid.  With  all  this  in  mind  it  is  not  astonishing  that 
two-thirds  of  our  cases  of  thyroid  abnormalities  were 
found  in  the  order  Camivora. 

The  anatomical  changes  of  the  human  thyroid  that 
precede  or  accompany  the  various  clinical  pictures  cannot 
be  said  to  be  uniform  to  a  degree  that  one  can  even 
approximately  predict  in  every  case  what  will  be  found 
at  operation,  at  autopsy  or  by  the  microscope.  Further- 
more, much  discussion  has  existed  upon  the  importance 
of  the  several  changes,  the  association  with  clinical  phe- 
nomena and  the  nomenclature.  I  shall  not  enter  the 
academic  discussion  with  our  material  because  so  much 
has  depended,  in  human  medicine,  upon  symptoms,  signs 
and  chemistry — data  that  we  cannot  adduce.  Upon  many 
occasions  I  have  seen  animals  with  very  evident  goitres, 
but  have  not  been  able  to  detect  bulging  of  the  eyes  or 
especial  nervousness.  One  striped  hyena  carried  his 
mass  for  several  years.  It  swelled  up  occasionally  and 
seemed  to  cause  dyspnoea.  At  one  examination  of  the 
beast,  to  see  if  anything  could  be  done  for  him,  a  large 
cyst  broke  under  the  examining  hand,  whereupon  a  deep 
inspiration  was  heard  and  reUef  was  apparent.  Tliis 
handling  was  repeated  twice,  these  times  wdth  the  pur- 
pose of  breaking  cysts  and  when  this  w^as   successful 


THE  DUCTLESS  GLANDS  323 

disappearance  of  the  dyspnoea  was  observed.  However, 
a  similar  attempt  upon  another  hyena  and  a  wolf  failed 
possibly  because  no  large  thin-walled  cyst  was  present. 
These  and  the  case  of  the  lion  cub  (page  170)  are  the 
only  instances  in  which  the  enlarged  thyroid  seemed  to 
have  given  serious  difficulty,  and  the  symptoms  were 
probably  due  to  pressure.  Many,  indeed  most,  enlarged 
thyroids  have  been  found  at  autopsy,  when  the  Garden 
personnel  was  unaware  of  their  existence.  Interesting 
notes  of  familial  cretinism  will  be  found  under  the 
appropriate  heading. 

Pathological  Anatomy.    Classification. 

And  now  to  return  to  the  question  of  morbid  anatomy 
of  the  thyroid  gland,  I  shall  begin  by  outlining  briefly  the 
classification  to  be  used  in  analyzing  our  cases,  a  system 
which  combines  those  of  many  pathologists,  yet  which  I 
believe  contains  the  essentials  of  all.  The  changes  in  the 
gland  being  hyperplastic  and  recessive,  at  times  to  a  stage 
of  atrophy,  no  clear .  cut  definite  line  of  demarcation 
separates  all  these  pictures ;  instead  they  must  be  thought 
of  as  merging  into  one  another.  When  the  thyroid 
enlarges  more  or  less  continuously  with  a  maintenance 
of  considerable  colloid,  the  picture  is  that  of  colloid 
goitre.  The  gland  is  pale,  gelatinous,  tense  but  resilient 
and  may  show  large  cystic  areas  with  fluid  contents. 
Microscopically  studied  the  acini  are  overfilled  with  col- 
loid yet  the  lining  cells  are  retained  but  flattened.  The 
cysts  may  show  the  ruptured  septa  of  the  original  acini. 
Enlarged  soft  reddish  glands  are  found  at  times  to  con- 
tain much  colloid,  nearly  every  acinus  being  distended 
with  it,  but  in  such  organs  the  epithelia  are  high  or  even 
reduplicated;  the  amount  of  contents  is  the  striking 
feature.  These  are  termed  hyperplasia  with  colloid. 
Hyperplasia  may  go  on  with  the  absorption  of  colloid, 
hyperplasia  without  colloid.  The  gland  is  then  a  darker 
body  of  more  solid  character,  red,  dull  purple  or  uni- 


324   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

formly  pale  pink,  somewhat  dependent  upon  the  blood 
content.  By  magnification  one  sees  smaller  acini  wdth 
prominent  high  cuboidal  or  cylindrical  epithelia  and  little 
or  no  colloid.  The  increase  of  cells  seems  to  be  due  both 
to  an  increase  in  their  size  and  number.  As  the  pro- 
liferation increases  the  lining  layer  must  be  accommo- 
dated so  that  it  bulges  out  into  the  lumen  as  a  bud  or 
papilla  which,  if  it  be  extensive  or  universal  in  the 
thyroid,  gives  rise  to  the  ADENOMATom  goitre  of  papil- 
lomatous TYPE.  This  growth  is  accompanied  by  much 
congestion  and  small  or  large  hemorrhages  may  occur, 
forming  cysts  containing  a  blood-stained  fluid.  Grossly 
such  a  gland  has  solid  and  cystic  areas,  is  mottled  red  and 
gray  or  brown  from  old  pigmentation  and  is  usually  of 
very  irregular  shape.  These  forms  are  more  or  less 
uniform  and  general,  but  in  certain  instances  the  hyper- 
plasia tends  to  remain  in  isolated  areas  or  nodules,  and 
in  these  develop  solid  masses  of  thyroid  epithelium,  some- 
times with  a  small  lumen  usually  devoid  of  colloid,  and  a 
rather  rich  but  loose  fibrous  supporting  tissue,  the  whole 
picture  resembling  the  microanatomy  of  the  fetal  gland ; 
to  these  the  name  fetal  adenoma  has  been  given,  but  since 
they  are  not  fetal  in  origin  and  do  not  behave  like  tumors 
I  have  called  them  nodular  adenomatoid  hyperplasias. 
The  next  step  in  hyperplasia  would  remove  it  from 
benign  to  malignant  in  pathological  character,  and  the 
term  neoplastic  hyperplasia  is  used ;  this  must  of  course 
be  limited  to  the  epithelial  growths,  since  sarcomata, 
while  they  occur  in  the  gland,  come  from  other  cells. 

The  changes  in  atrophy  consist  in  irregular  distortion 
of  the  gland  by  fibrous  tissue  to  which  may  be  added  large 
colloid  or  fluid  cysts.  There  is  no  uniform  finding  in  the 
thyroid  for  the  diseases  believed  to  be  due  to  its  atrophy, 
functionally  at  least,  namely  cretinism  and  myxedema. 
In  the  former  there  may  be  no  thyroid,  or  it  may  rather 
closely  resemble  the  normal  organ  while  in  the  latter 
definite  scarring  and  distortion  is  the  rule.    The  gland 


THE  DUCTLESS  GLANDS  325 

aciiii  are  compressed,  the  cells  vacuolated  or  crushed  out 
of  existence  or  there  may  be  colloid  cysts. 

Inflammations  occur  as  swellings  of  the  interstitial 
tissues  and  of  the  acinus  cells  during  many  acute  infec- 
tions. Repetitions  of  this  may  leave  a  definite  increase 
of  connective  tissue  with  large  cells  in  the  acini,  a  lesion 
which  many  observers  have  looked  upon  as  underlying 
certain  goitres  and  myxedema. 

Hypeeplasias. 

The  cause  of  progressive  hyperplasias  has  been 
ascribed  to  infection,  to  chemical  substances  in  water  and 
food,  endogenous  toxins,  heredity  and  many  other  fac- 
tors. While  we  can  add  nothing  definite  in  this  matter  it 
is  worthy  of  notice  that  all  our  animals  are  exposed  to  the 
same  general  climatic  conditions,  receive  the  same  water, 
are  fed  from  the  same  stocks  and  many  varieties  may  be 
in  charge  of  the  same  keeper.  The  influence  of  preexisting 
infections  cannot  of  course  be  measured.  Inbreeding  or 
captive  breeding  seems  to  have  a  very  definite  effect  upon 
thyroid  insufficiency  as  is  well  known  and  so  sharply 
emphasized  by  McCarrison  in  his  reference  to  intermar- 
riage among  certain  Moslems ;  I  shall  cite  the  history  of 
a  wolf  bitch  which  gave  birth  to  three  cretin  litters  while 
apparently  well  but  mated  to  a  goitrous  male.  These  facts 
concerning  the  etiology  are  given  merely  to  emphasize  the 
high  degree  of  probability  that  the  distribution  of  the 
lesions  of  the  thyroid  gland  among  our  specimens  indi- 
cates the  susceptibility  of  the  various  orders.  This  per- 
haps needs  no  emphasis  for  the  carnivores,  but  it  does  for 
the  marsupials.  The  literature  contains  many  references 
to  goitre  in  domesticated  ungulates ;  this  would  give  the 
impression  that  they  are  common  among  them,  and  so 
they  may  be,  but  this  is  not  the  case  for  wild  ungulates. 
There  being  no  doubt  that  the  Carnivora  have  the  highest 
incidence  of  thyroid  enlargement,  man  being  especially 
prone  to  it,  and  since  goitre  may  be  induced  in  fish  by 


326   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 


feeding  meat,  the  inference  is  direct  that  high  protein 
diet  stands  in  some  relation  to  this  condition.  Thirty- 
nine  of  our  sixty  thyroid  lesions  occurred  in  the  order 
Carnivora;  all  the  families  of  land  varieties  are  repre- 
sented ;  8.1  per  cent,  of  the  specimens  coming  to  autopsy 
showed  definite  thyroid  alterations.  However,  marsu- 
pials have  also  a  decided  thyroid  vulnerability  as 
indicated  by  4  per  cent,  of  the  specimens  presenting 
abnormalities  at  death.  Four  of  the  seven  cases  were 
among  the  carnivorous  opossums  and  ''devils,"  the 
remaining  three  being  in  herbivorous  kangaroos.  The 
influence  of  high  protein  diet  is  not  evident  in  birds. 

A  discussion  of  the  essential  pathology  can  be  based 
upon  the  accompanying  table.  In  making  a  diagnosis 
care  was  used  to  exclude  mild  swelling  of  the  gland  seen 
in  acute  infectious  disease  and  under  conditions  of  sexual 
activity.  The  gross  diagnosis  was  checked  by  micro- 
scopical section,  and  all  but  a  very  few  have  been 
reexamined  for  the  purpose  of  making  the  table. 

Table  16. 
Shoxving  Distribution  of  Lesions  in  the  Thyroid  Body  by  Giving  the  Number  of 
Cases  Met  in  the  Autopsies  upon  the  Various  Orders,  According  to  the  Classifi- 
cation Given  in  the  Text. 


A  ,1 

R 

2 

X 

JS 

il 

E 

2 

V 

» 

» 

^ia 

a  03 

c« 

2 

Order 

Is 

•5 
O 

T3 
O 

ll 
13 

•5 

II 
13 

^1 

il 

.1 

a  a 
catU 

1 
1 

1 

1 

O 

o 

K 

K 

^ 

'^ 

^ 

s 

w 

< 

Carnivora 

89 

7 

6 

5 

4 

Q 

1 

1 

1 

7 

Rodentia 

1 

1 

Ungulata 

2 

1 

1 

Marsupialia . 

7 

1 

1 

1 

1 

s 

2 
2 
1 

2 

1 

1 

1 

Psittaci 

Gain 

2 

1 

1 

1 

1 

Alectorides 

Anseres 

3 

2 

Total 

60 

12 

11 

7 

6 

12 

2 

1 

2 

8 

Fig.    31.— hyperplasia  with  colloid.     AMERICAN    BADGER  (TAXl- 
DEA  TAXUS).     MEASUREMENTS.  3  X  1.5  CM.;    3  X  1.5  X  1^  CM. 


J'% 


Fig.   32.— adenomatoid   GOITRE.       RACCOON-I.IKE   DOG   (CANIS   PROCYONOII 


THE  DUCTLESS  GLANDS  327 

This  rather  diversified  group  of  pathological  lesions 
would  warrant  one  to  expect  a  notable  number  of 
instances  of  disease  observed  during  life,  suggesting  that 
the  thyroid  was  at  fault ;  such,  however,  is  not  the  case. 
In  the  first  place,  no  case  of  exophthalmic  goitre,  as  the 
symptom  complex  is  known  in  man,  has  been  observed,  yet 
the  anatomical  alterations,  hyperplasia  without  colloid, 
and  with  papillary  or  solid  adenomatoid  character,  are 
abundantly  represented.  For  the  pathologist  to  accept 
a  case  as  toxic  goitre  I  would  ask  evidence  of  enlargement 
of  the  heart  and  perhaps  in  addition  degeneration  of  the 
myocardium.  The  animal  showing  the  closest  resem- 
blance to  the  disease  in  man  was  a  Raccoon-like  Dog 
{Canis  procyonoides)  whose  history  and  notes  are  given 
in  brief. 

Raccoon-like  dog  {Canis  procyonoides)  9.  Acute  hemorrhagic 
splenitis.  Acute  fermentative  gastritis.  Subacute  catarrhal  enteritis. 
Acute  general  infection.  Hypertrophic  cirrhosis  of  liver.  Chronic 
interstitial  nephritis.  Hypertrophy  of  heart  with  acute  myocarditis- 
infiltrative  and  parenchymatous.  Chronic  lymphadenitis  with  acute 
exacerbation.  Epigastritic  and  gastric  venous  stasis.  Nodular  adenoma- 
tous goitre.  The  right  thyroid  is  lower,  both  measure  4x3x2  cm.,  are 
soft,  resilient  with  dense  gray  capsule.  Section  shows  cysts  filled  with 
blood  separated  by  pale  septa  of  soft  tissue  of  varying  thickness.  One 
similar  mass  under  angle  of  jaw  seemed  like  a  lymph  node  but  on  sec- 
tion is  like  thyroid.  Parathyroids  not  found.  Pleurae  negative.  Lungs 
are  gray,  collapsed  except  lower  half  of  lower  lobes  which  are  slightly 
emphysematous  and  edematous.  No  consolidations.  Pericardium  con- 
tains about  3  cc.  clear,  colorless  fluid.  Epicardium  is  glistening,  trans- 
parent and  pale.  Heart  muscle  is  soft  flaccid,  pale  mottled  gray-brown. 
All  chambers  are  distended  with  mixed  clot.  The  coronary  muscles  and 
columnas  are  mottled  brown  and  gray.  The  tips  at  insertion  of  chordae 
are  pale.  Streaks  of  gray  run  through  muscles.  One  area  1x2  cm. 
of  softening  found  in  middle  of  left  auricular  muscle.  Aorta  nega- 
tive. The  liver  is  enlarged,  surface  rough  and  irregular,  edges  rounded 
and  uneven,  consistency  firm  and  tough,  color  mottled  deep  red-brown. 
Section  surface  glistening,  moist,  granular,  opaque.  Lobular  markings 
not  lost  but  obscured.  Connective  tissue  lines  not  clear  but  surely 
diffuse  in  lobules.  Scars  on  surface  leading  to  definite  connective  tissue 
strands  about  vessels  and  irregularly  placed.  Gall-duct  patulous 
with  limpid  bile.  Spleen  is  well  forward  in  front  and  below  stomach. 
It  is  much  enlarged,  soft,  tough,  has  a  smooth,  tense  capsule.     Section 


328  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

surface  shoAvs  homogeneous  purple  pulp  Avith  faint,  narrow  but  tough 
trabeculae.  Follicles  distinctly  outlined,  slightly  large  but  merely  of 
a  slightly  paler  purple  than  pulp.  On  surface  are  many  round  2-5  mm. 
sharply  outlined  gray  thickenings  of  the  capsule  and  immediately  sub- 
jacent pulp.  The  right  kidney's  lower  half  has  been  replaced  by  a 
thin-walled  clear  cyst  3  x  2.5  x  2  em.  The  left  kidney  is  small,  capsule 
strips  with  difficulty  tearing  surface  slightly.  Surface  and  section  are 
mottled  pink  and  gray,  glistening  and  opaque.  It  is  firm,  dense  and 
tough,  cortex  narrow,  medulla  wide.  Cortex  has  obscure,  irregular  mark- 
ings with  few  small  cysts,  striae  and  glomeruli,  faintly  visible,  margin 
between  layers  iiTegular.  Veins  are  distended  over  surfaces  of  stomach, 
under  surface  of  diaphragm,  in  peritoneum  over  liver  but  not  in  ab- 
dominal Avail.  Stomach  contains  sour  gas  and  water.  Mucosa  especially 
near  cardia  is  deep  pui-ple.  Rugae  are  large  and  pennanent  but  mucosa 
and  submucosa  are  soft  and  on  section  congestion  does  not  extend  deep. 
The  tips  of  the  rugae  near  pylorus  are  infiltrated  and  the  infiltration  is 
surrounded  by  a  zone  of  congestion.  Mucosa  seems  about  to  slough 
but  has  not  separated.  At  pylorus  mucosa  becomes  deep  broAvn-red, 
dense,  swollen,  opaque  and  coA-ered  Avith  a  slimy,  soft  brown  mucus. 
Folds  are  prominent  but  temporary.  In  jejunum  and  ileum  mucosa  is 
still  swollen  and  opaque  and  rugae  are  still  larger  than  normal  and  tem- 
porary Avith  a  dense  sensation  on  compressing  them.  The  color  is  not 
broAvn  but  deep  pink  and  yellow  Avith  areas  of  submucous  injection. 
Follicles  not  visible.  Colon  is  negative  except  for  slight  thickening  of 
mucosa  unaccompanied  by  congestion  or  opacity.  Lymph  glands  of 
small  omentum  are  small,  firm,  yelloAV,  homogeneous;  those  of  the  mes- 
entery are  large,  edematous,  yellow  brown  and  tense  Avith  lymph  Avhieh 
escapes  on  section.  LjTnph  channels  up  to  mesenteric  stalk  can  be 
traced. 

Histological  Notes. — LiA'er  architecture  much  altered  by  passively 
dilated  hepatic  capillaries  chiefly  toward  the  centre  of  the  distorted 
lobules.  This  distortion  is  in  the  form  of  irregular  liver  columns  sepa- 
rated by  irregular  vessels  and  interlobular  connective  tissue.  This  latter 
is  increased  everywhere  but  is  abnormal  in  distribution  Avithin  the 
lobules.  The  connective  tissue  at  the  portal  spaces  is  not  so  much 
increased  but  it  shoAvs  most  around  arteries.  Bile  ducts  seem  not 
increased  in  numbers.  Much  bile  pigment  in  large,  coarse,  dark  brown 
masses  chiefly  settled  Avithin  portal  spaces.  The  cells  show  slight  fatty 
infiltration.  No  multinucleation.  Organ  is  not  seriously  robbed  of 
functionating  tissue.  Alterations  are  not  equally  distributed  OA'er 
section.  Spleen  shoAvs  enormous  congestion  with  edema  of  the  few 
chords  and  perifollicular  tissue  left  unengorged  Avith  blood.  Follicles 
are  negative.  No  connective  tissue  increase.  Blood  destruction  not 
noAv  actiA'e  but  there  are  many  hemosiderotic  masses  irregularly  scattered. 
The  subcapsular  areas  are  loose  edematous  follicles.  Heart  muscle 
fibres  have  lost  all  transverse  striations,  some  are  hyaline  Avhile  others 
are  fibrillar.     The  nuclei  are  decreased  in  number  but  there  is  no  in- 


THE  DUCTLESS  GLANDS  329 

crease  of  connective  tissue  nuclei.  No  pigmentation.  There  are  several 
areas  of  round  and  polynuclear  cell  infiltration  and  one  distinct  abscess 
in  section.  The  perivascular  tissues  are  edematous.  Muscle  fibres  are 
large  and  wide.  Thyroid  made  up  largely  of  slightly  enlarged  acini  in 
most  of  which  a  slightly  eosin-stained  hyaline  collection  is  found. 
There  are  a  few  cysts  containing  a  thrombus  and  hemorrhage.  There 
are  no  typical  colloid  cysts.  Some  scars  from  old  hemorrhages  may  be 
seen.  There  is  much  free  blood  in  and  between  acini.  Blood  pigment 
free  and  in  granule  cells  is  abundant.  Some  acinus  cells  show  fat  drop- 
lets. Lung  shows  old  interstitial  tissue  increase  especially  about  vessels 
and  a  few  scars,  some  of  Avhich  are  forming  cartilage.  These  are  deeply 
encapsulated.  Mesenteric  lymph  nodes  show  trabecular  thickening  with 
active  connective  tissue  formation  which  is  also  present  about  follicles 
and  along  edges  of  chords.  Follicles  lack  germ  centres,  solidly  lymphatic. 
About  them  and  in  and  along  chords  are  many  tissue  ceils  some  of 
Maximov  type  and  a  few  eosinophiles.  Many  of  these  and  endothelial 
cells  are  phagocytic  of  red  blood  cells.     (Fig.  32.) 

Just  at  the  time  of  completing  this  book  another  case 
strongly  resembling  exophthalmic  goitre  in  man  was 
encountered  in  a  Gray  wolf  {Canis  lupus).  This  animal 
had  a  history  of  enlarged  neck  and  enlarging  abdomen  for 
about  six  months.  His  appetite  and  discharges  remained 
about  normal  but  weight  was  lost  and  activity  reduced  to 
a  minimum.  Attempts  at  removing  the  fluid  believed  to 
be  in  the  peritoneum,  by  the  use  of  diuretics,  failing  and 
the  beast  being  in  such  poor  shape,  he  was  killed.  An 
enormous  adenomatoid  goitre,  concentric  hypertrophy  of 
the  heart,  passive  dilatation  of  all  cervical  and  thoracic 
veins,  passive  congestion  of  the  liver  and  congestion  of 
the  portal  area  were  autopsy  diagnoses.  It  will  be 
noted  that  no  exophthalmos  and  nervousness  were 
observed  during  life. 

Bone  disease  and  atheroma  are  at  times  associated 
with  thyroid  insufficiency  in  man.  The  former,  aside  from 
osteogenesis  imperfecta  of  cretinism,  occurred  only  once 
in  a  carnivore  and  once  in  a  marsupial.  There  is  but 
one  case  of  atheroma  among  the  sixty  cases  of  thy- 
roid disease. 

The  reaction  of  the  avian  thyroid  in  its  hyperplasias  is 
somewhat  different  from  that  of  the  mammalian.     The 

22 


330   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

delicacy  of  the  septa  and  the  relative  paucity  of  vessels  is 
perhaps  the  reason  that  the  gross  and  microscopic  pic- 
tures differ  from,  those  found  in  mammals.  It  should  be 
emphasized,  however,  that  while  one  can  perceive  a  hyper- 
plasia of  the  gland  of  both  classes  when  the  testes  or 
ovaries  are  active,  there  appears  less  participation  of  the 
thyroid  in  birds  in  infectious  diseases  than  is  the  case  for 
mammals.  In  simple  functional  hyperplasia  the  capsu- 
lar vessels  are  prominent,  but  the  cross  section  need  show 
no  change.  In  the  continued  hyperplasias  the  organ 
remains  more  solid,  being  less  apt  to  develop  cysts ;  large 
cysts  are  occasionally  seen,  however,  and  in  one  case  the 
entire  gland  was  composed  of  them.  Microscopically  the 
differences  are  largely  of  degree  in  that  the  process  is  less 
frank  in  development,  but  the  essential  changes  of  swollen 
epithelia  and  condensed  colloid  remain  the  same. 

Atrophies. 

The  thyroid  gland  in  its  functional  capacity,  may  be 
considered  to  undergo  hyperplasia  and  then  atrophy  of 
the  parenchyma  cells.  Normally  this  would  leave  the  col- 
loid, the  epithelia  and  the  supporting  tissue  in  proper 
balance,  but  in  the  presence  of  low  grade  inflammation  or 
where  an  abnormally  hyperplastic  process  retrogresses, 
the  connective  tissues  may  exceed  their  norm,  the 
epithelia  may  be  shed  or  remain  high  and  the  colloid  be 
irregular  in  distribution.  Such  a  state  of  atrophy  may 
exist  in  fetal  life,  arise  from  unknown  cause  during  a 
course  of  toxic  goitre,  or  perhaps  insidiously  in  chronic 
toxic  conditions.  When  this  occurs  in  fetal  life  cretinism 
or  myxedema  arises,  when  in  later  life,  only  the  latter 
appears.  Judged  entirely  by  microscopic  findings,  eight 
instances  of  atrophic  changes  in  the  thyroid  have  been 
found.  Three  of  these  were  in  Carnivora  and  were 
secondary  to  definite  goitres,  but  were  not  followed  by 
myxedema ;  one  of  these  three  was  a  cretin.  A  brother 
of  this  cretin  but  not  himself  a  cretin,  died  at  the  age  of 


Fig.  33. — COLLOID  GOITRE  WITH  HEMORRHAGE  FROM  LEFT 
GLAND  WHICH  KILLED  THE  BIRD.  BLACK  AREA  IS  CLOTTED 
BLOOD.      MUTE  SWAN   (CYGNUS  OLOR6  ). 


Fig.  54. — AT  THE  LEFT,  THE  INSIDE  OF  THE  CAI.VARILM 
SHOWING  HEMORRHAGIC  PACHYMENINGITIS.  AT  THE  RIGHT  IS 
A  FEMUR.  SHORT  AND  HEAVY  BUT  WITH  THIN  CORTEX;  CONSTRUC- 
TION IS  ORDERLY.      CRETIN  GRAY  WOLF  PUP  (CANIS  MEXICANUS). 


THE  DUCTLESS  GLANDS  331 

five  months  from  acute  dilatation  of  the  heart,  and  with  a 
decidedly  atrophic  thyroid  gland.  A  lion  showed  a  dis- 
torted gland,  the  result  of  chronic  inflammation,  a 
condition  also  present  in  a  leopard,  in  the  latter  possibly 
in  association  with  a  general  infection  of  the  heart,  ves- 
sels and  kidneys  of  long  standing.  A  case  in  a  bear  can 
only  be  explained  on  the  basis  of  chronic  intestinal 
toxemia.  A  camel  is  the  only  other  variety  of  animal  to 
show  this  regressive  change.  The  beast  suffered  with  a 
marked  anemia  with  marrow  atrophy  and  hydatid 
disease ;  calcification  was  found  in  the  thyroid. 

As  has  been  repeatedly  stated,  myxedema  has  not  been 
seen,  but  hypothyroidism  has  expressed  itself  in  these 
animals  as  cretinism.  The  most  interesting  pathological 
fact  concerning  the  relation  of  the  thyroid  to  this  mal- 
development  is  that  there  is  absolutely  no  uniform  gross 
or  microscopical  change  constantly  present  in  the  typical 
cases.  This  will  become  more  evident  as  the  following 
records  of  our  cases  are  perused.  In  1914  an  apparently 
normal  Gray  Wolf  bitch  threw  two  normal  young  ones 
which  died  of  lack  of  maternal  care ;  they  were  not  posted. 
The  father  of  this  litter  died  shortly  and  was  found  to 
have  a  sarcomatous  hyperplasia  of  the  thyroid.  A  year 
after  the  first  lot  a  second  litter  was  born  of  an  appar- 
ently perfect  father  which  still  lives.  This  animal  was 
purchased  in  the  same  lot  with  the  mother,  and  the  two 
could  be  related.  The  first  father  was  not  related  to  the 
female.  This  litter  consisted  of  seven,  two  dying  almost 
at  once  and  burned,  the  other  five  not  being  especially 
good  specimens.  They  died  at  ages  ranging  from  two  to 
five  months  and  were  all  cretins  or  cretinoid.  Two  showed 
hemorrhagic  pachymeningitis,  one  external,  one  internal, 
and  the  usual  bulky  skeleton  of  cretinism  (Fig.  34). 
The  bones  were  constructed  in  a  rather  orderly  and  some- 
what graceful  manner,  the  uncalcified  epiphyses  being 
only  occasionally  distorted.  The  same  two  animals, 
mated  again  in  1916,  had  as  offspring  seven  pups.    One 


332  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

evident  cretin  was  killed  while  another  runt  was  sacrificed 
and  found  to  have  fractures  of  both  femora  around  Avhich 
no  trace  of  callus  was  discoverable  (Fig.  35).  Two  other 
cubs  were  apparently  normal,  while  the  remaining  three 
did  not  develop  and  soon  showed  the  cretin  characters. 
When  this  group  was  about  three  months  old  they  were 
fed  chopped  horse  thyroid;  one  improved  decidedly,  one 
slightly,  the  third  not  at  all,  but  it  might  have  been  too 
weak  to  get  its  share.  These  animals  lived  from  eight 
months  to  three  years ;  the  two  good  ones  remain  alive. 
In  1919  the  mother  was  killed  by  her  cubs,  probably 
because  she  was  weakened  by  long  sickness.  A  papillary 
adenomatoid  goitre,  endocarditis,  nephritis  and  chronic 
enteritis  were  found.  The  pathological  changes  in  the 
thyroids  of  the  cretins  were  as  follows:  In  the  second 
litter  two  cubs  had  hyperplasia  with  colloid,  one  had 
nodular  adenomatoid  change  and  the  fourth  showed  dis- 
tinct atrophy  secondary  to  colloid  increase.  In  the  third 
group  two  had  distinct  colloid  changes,  once  pure  and 
once  as  a  secondary  process  with  some  evidence  of 
atrophy  to  alter  the  fibrous  tissue  and  shape  of  the  acini. 
The  remaining  four  seem  to  be  all  colloid  in  character,  but 
I  am  not  satisfied  with  the  description  or  sections  so  that 
I  shall  not  offer  an  unqualified  diagnosis.  The  adrenals 
of  these  animals  all  showed  some  medullary  congestion 
but  no  change  in  the  chromaffin  or  lipoidal  content. 

The  deformative  lesions  of  bones  are  frequently  asso- 
ciated mth  lesions  in  the  pituitary  body.  Several  of  our 
thyroid  cases  have  been  studied  for  such  changes  without 
their  discovery.  Indeed  no  gross  alterations  have  been 
noted  among  many  hundreds  of  hypophyses  seen  in 
removing  the  brain  nor  in  a  small  number  studied 
histologically.  Those  examined  under  the  microscope 
have  seemed  to  correspond  to  the  descriptions  given  by 
Stendell  in  Oppel's  Handbook  of  Comparative  Micro- 
scopic Anatomy. 


kk;.  55.— pathological  kracturk  of  fk.vr'r.  cretin  gray  wolf  pup. 


THE  DUCTLESS  GLANDS  333 

Tumors. 

True  malignant  hyperplasias  of  the  thyroid  epithelium 
in  man  are  being  more  thoroughly  studied  in  recent  times 
so  that  similar  lesions  in  the  lower  animals  gain  interest. 
The  notes  given  below  are  of  value  as  individual  observa- 
tions only,  but  since  three  were  in  carnivores,  another 
indication  is  at  hand  of  the  vulnerability  of  this  order. 
One  of  the  cases  is  admitted  upon  diagnosis  alone,  the 
slides  and  records  having  been  lost,  but  since  the  deter- 
mination was  made  by  Dr.  C.  Y.  White,  I  am  satisfied  to 
accept  it.  The  four  of  which  notes  are  at  hand  are 
as  follows : 

Raccoon-like  Dog  {Canis  procyonoides)  6.  Adenocarcinoma  sarco- 
matodes. Metastases  to  liver  and  lungs.  Fatty  degeneration  of  liver 
and  kidney.  Acute  diffuse  splenitis.  Submucous  hemorrhages  in 
stomach.  At  level  of  thyroid  cartilage  on  each  side  and  removed  1  cm. 
from  same  is  a  rounded  encapsulated  nodule  measuring  2.5  x  2  x  1  cm. 
Portions  are  hard,  others  fluctuate  suggesting  cystic  degeneration.  Below 
these  nodules  are  two  bodies  also  bilateral,  evidently  lobes  of  thyroid, 
each  measuring  5  x  2.5  x  2.5  cm.  They  are  firm  with  some  foci  of 
cystic  softening.  From  a  ruptured  cyst  of  the  right  lobe  grumous,  red, 
malodorous  material  exudes.  Peripheries  of  such  cyst  show  greenish 
discoloration.  Bodies  as  a  whole  are  greenish  black  in  color.  They  are 
well  encapsulated,  do  not  meet  in  midline  but  are  joined  at  lower  pole 
by  firm,  apparently  colloid,  material.  All  through  lung  especially  under 
pleura  there  are  dark  red,  rounded,  firm,  well  circumscribed  foci  measur- 
ing 2-8  mm.  diameter.  They  project  markedly  on  pleural  surface.  No 
capsule  can  be  made  out.  Upon  incising  they  have  lighter  red  centres 
and  deeper  peripheries.  They  cut  with  resistance  and  have  no  inclination 
toward  a  wedge  shape.  Surface  is  for  most  part  smooth  except  where 
tumors  are  present.  Organ  is  soft  and  distinctly  yellow.  All  portions 
of  liver  contain  rounded  and  irregular  nodules,  some  deep,  others  super- 
ficial. They  vary  in  size  from  2  mm.  to  3.5  cm.  diameter.  No  capsule 
can  be  made  out,  yet  they  are  circumscribed.  The  central  portions  of 
larger  nodules  are  dirty  gray  and  friable.  Peripheral  parts  dark  red. 
Smaller  lesions  are  solid  red  and  of  fleshy  consistency.  Spleen  is  deep 
dark  red,  homogeneous.  Histological  section  of  thyroid  shows  firm, 
old,  dense  capsule  very  irregular  in  thickness  seemingly  on  account  of 
the  penetration  of  the  enclosed  tumor  cells.  Such  infiltration  gives  the 
inner  outline  of  capsule  a  very  irregular,  bizarre  appearance,  and  at 
times  thins  the  capsule  until  it  is  reduced  to  nil.  In  one  place  the 
tumor  elements  appear  outside  the  capsule  at  a  point  where  a  large 
vessel  is  apposed  to  outside  capsule.   The  appearances  within  this  cap- 


334   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

sule  vary;  in  places  the  picture  is  that  of  a  carcinoma.  Small,  round, 
interspersed  wdth  larger  irregularly  shaped  acini  are  seen  lined  by  a 
single  layer  of  low  cuboidal  epithelium.  \ery  frequently  indeed  the 
lining  cells  contain  fine  granules  of  golden  brown  pigment  even  where 
their  lumina  contain  no  blood.  Some  of  the  larger  acini  contain  altered 
blood  cells  and  a  smooth,  pink  material,  knife  streaked  and  vacuolated 
peripherally.  In  some  parts  of  section  these  acini  are  regular  and 
well  formed,  in  others  they  are  very  irregular  and  appear  to  be  eroding 
the  capsule.  A  second  appearance  concerns  the  connective  tissue.  Ap- 
pearing in  almost  any  part  of  the  section  and  bearing  no  regular  rela- 
tion to  the  epithelial  elements  or  the  section  in  general  are  areas  of 
closely  packed  large  spindle  cells  with  hyperchromatic  nuclei.  In 
another  place  such  spindle  cells  are  arranged  purposefully  to  form 
irregular  capillaries  containing  blood.  A  third  appearance  results  from 
a  combination  of  the  first  two.  Here  there  are  acini,  between  which 
run  blood  capillaries  with  remarkably  rich  and  numerous  embi'yonic 
lining  cells.  A  section  stained  by  Van  Gieson  stain  proves  that  part 
of  the  pink  intra-alveolar  material  is  colloid.  Every  gradation  can  be 
made  out  in  tint  of  this  material  from  pink  to  salmon  to  orange.  It 
is  often  veiy  difficult  or  impossible  to  state  whether  a  given  blood- 
filled  space  is  a  blood  vessel  or  an  acinus  with  hemorrhage.  In  both 
structures  the  lining  consists  of  flattened  cells.  In  one  there  is  the 
possibility  of  colloid,  in  the  other  of  hematogeneous  hyaline,  both 
with  peripheral  vacuolization.  Lung  shows  walls  of  alveoli  thickened 
by  young  type  of  cells.  Nuclei  of  cells  lining  bronchi  are  prominent, 
in  good  condition.  Much  coal  pigment  through  whole  section.  Air 
sacs  empty.  There  are  several  rounded  nodes  through  section  con- 
sisting of  closely  packed  spindle  and  round  cells.  Blood  is  abundant 
in  such  nodules  both  in  small  lined  spaces  like  capillaries  and  in  larger 
necrotic  foci  where  there  is  abundant  blood  pigment.  In  one  place 
an  irregrular,  large  acinus  is  seen  containing  a  smooth  pink  material. 
A  large  part  of  the  interstitial  tissue  is  diffusely  infiltrated  by  the  large, 
round  cells  with  hyperchromatic  nuclei  (Figs.  33  and  37). 

Prairie  Wolf  {Canis  latrans)  6.  Mixed  tumor  of  thyroid.  Metas- 
tases to  lungs.  The  neck  of  the  animal  is  enormously  enlarged,  the 
diameter  exceeding  that  of  the  body.  Thyi'oid  is  enormously  enlarged 
to  about  the  size  of  a  child's  head,  rather  firm  before  incision.  When 
incised  about  300  cc.  blood  stained  fluid  drained.  It  is  rather  soft 
and  quite  friable  looking  as  if  made  up  of  fatty  and  hemorrhagic 
matter.  The  lung  is  of  mottled  deep  red  color  with  here  and  there  on 
surface  small  hemispherical  areas  about  the  color  of  the  surrounding 
tissue  but  of  slightly  increased  resistance.  They  are  raised  above  the 
surface  and  measure  2-7  mm.  in  diameter.  Histological  section 
of  thyroid  shows  a  mixed  tumor.  It  is  not  possible  to  say  that  it  is  a 
pure  thjToid  gland  tumor.  It  is  largely  sarcomatous,  the  round  cell 
alveolar  arrangement  dominant  at  one  place,  at  another  the  short 
spindle  cell  but  not  typical,  so-called  spindle  celled  type.     There  are 


THE  DUCTLESS  GLANDS  335 

many  areas  of  small  and  a  few  of  large  hemorrhage.  Cartilaginous 
deposit  is  occurring  at  some  places  in  the  field  showing  the  latter  type 
of  sarcoma.  Section  of  lung  contains  a  large  sarcoma  nodule.  The 
cells  consist  of  round  cells  without  the  large  cartilage-like  cells  found 
in  the  original  tumor  mass. 

Coypu  (Myocastor  coypus)  6.  Sarcoma  of  right  thyroid.  The 
thyroids  occupy  a  position  deep  in  the  neck  upon  the  anterior  vertebral 
muscles,  the  left  higher  than  right,  being  up  to  level  of  top  of  thyroid 
cartilage.  Only  a  half-inch  of  lower  pole  of  right  remains  and  it  is 
like  the  left  which  is  soft,  deep  brown-red,  delicately  lobulated,  closely 
bound  to  trachea  but  movable  in  fascia.  It  is  30  x  10  x  3  mm.  The 
upper  part  of  the  right  organ  is  occupied  by,  or  at  least  within  the 
same  capsule  as  a  25  x  15  x  10  mm.  encapsulated,  pink  mass  with  many 
small  vessels  on  its  exterior.  It  is  soft  and  on  section  the  surface  is 
mushy,  of  gray-pink-yellow,  and  seems  to  have  an  exceedingly  delicate 
trabecular  network.  Posterior  and  superior  to  this,  lying  near  the 
salivary  glands  but  back  of  them  is  a  similar  mass  15  x  12  x  5  mm.  Still 
another  lies  anterior  to  what  remains  of  the  right  thyroid  and  is  about 
8x5x4  mm.  The  adrenals  are  30  x  13  x  8  mm.  slightly  hard  and  not 
unlike  a  long  kidney  in  arrangement.  The  cortex  is  wide,  regular, 
brown  or  tawny,  the  medulla  rich  in  vessels  and  deep  brown.  His- 
tological section  of  thyroid  is  an  almost  completely  cellular  mass  with 
here  and  there  delicate  and  incomplete  trabeculations.  Small  blood 
vessels  are  numerous  and  consist  of  a  delicate  line  with  a  cell  nucleus 
here  and  there,  that  is  no  true  wall.  It  seems  as  if  the  blood  channels 
jvere  simply  regular  spaces  through  the  cell  mass.  The  cell  type  is 
mononuclear  with  definitely  acidophilic  "  granuloid  "  somewhat  vacuo- 
lated protoplasm.  The  nucleus  is  almost  without  exception  eccentric, 
rather  poor  in  chromatin  but  in  places  diffusely  staining.  Mostly, 
however,  the  nuclear  skein  is  in  spots  or  threads  and  fairly  dense 
around  margins,  therefore  not  unlike  a  thyroid  cell  and  a  plasma  cell. 
Here  and  there  one  finds  compressed  remains  of  thyroid  acini.  There 
is  decided  irregularity  of  size  and  shape  in  these  cells.  Its  origin  is  not 
clear  but  this  seems  like  a  sarcoma  of  the  thyroid. 

Undulated  Grass  Parrakeet  {Melopsittacus  undulatus)  6.  Medul- 
lary carcinoma  of  thyroid.  On  opening  the  body  a  mass  10  x  6  x  4  mm. 
is  found  in  the  upper  thoracic  region  on  the  right  side.  A  similar 
mass  measuring  5x3x2  mm.  lies  in  similar  position  on  the  left  side. 
They  are  identified  as  thyroid  glands  only  from  their  position  and 
from  the  numerous  large  vessels  which  radiate  from  them.  An  especially 
large  vessel  leads  directly  to  the  heart.  Long  axes  extend  anteroposteri- 
orly.  The  masses  are  of  a  firm  gelatinous  consistency,  the  color  of  carpen- 
ters glue  in  lower  portions,  shading  off  to  a  dirty  canary  yellow  above. 
They  have  a  translucent  appearance  in  lower  portions.  The  surface 
is  fairly  smooth,  adherent  latterly  and  posteriorly.  They  are  well 
circumscribed.     At  one  end  of  the  histological  section  thyroid  tissue 


336  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

is  easily  identified.  It  varies  from  normal  in  that  its  spaces  are  often 
very  large,  contain  villus  projections  or  may  be  completely  filled  by 
large  compound  granule  cells  with  no  colloid.  Other  acini  are  atypical 
and  contain  typical  colloid.  Continuous  to  such  thyroid  tissue  is  a 
very  large,  rounded  tumor.  It  consists  of  round  cells  with  round 
nuclei  in  which  many  mitotic  figures  may  be  seen.  An  arrangement 
into  acini  cannot  be  made  out  nor  is  colloid  material  abundant.  In 
one  or  two  places  an  irregular  collection  of  such  material  may  be  seen 
with  peripheral  vacuolization  but  its  confines  are  always  indefinite. 
As  far  as  section  goes  the  mass  is  well  encapsulated  but  the  lymphatics 
are  infiltrated  by  the  tumor  cells.  The  tumor,  too,  is  sharply  separated 
from  the  relatively  normal  thyroid.  Irregularly  scattered  through 
section  are  remarkable  cells  with  nuclei  three  or  four  times  the  size 
of  other  nuclei.  They  may  be  hyperchromatic  or  normally  staining. 
(Figs.  38  and  39.) 

The  THYMUS  BODY  is  a  structure  encountered  in  our 
specimens  with  greater  regularity  than  is  the  case  in 
human  autopsy  experience.  However,  no  great  size  of  the 
gland  is  observed,  and  there  is  no  record  or  recollection  of 
anything  which  could  resemble  an  enlargement  suggest- 
ing status  thymicolymphaticus  nor  has  a  tumor  with  this 
organ  as  its  origin  been  observed.  In  one  case  only  did 
the  thymus  present  what  was  believed  to  be  an  unusual 
size.  An  adult  Gray  Lagothrix  {Lagothrix  lagotricha) 
died  mth  an  acute  intestinal  infection.  Its  thymus  was  a 
large,  soft,  deep  pink  body  lying  in  the  anterior  mediasti- 
num, running  up  to  the  clavicular  joints  and  down  along 
the  sternum.  The  death  had  ample  explanation  without 
any  state  of  this  organ.  The  thymus  body  has  not  been 
found  enlarged  in  association  with  thyroid  disease. 

The  SUPRARENAL,  or  ADRENAL  BODY  is  an  orgau  of  essen- 
tially the  same  general  construction  in  the  two  classes 
here  studied  except  that  in  birds  the  cortical  portion  may 
be  imperfectly  developed  and  in  some  of  the  lower  groups 
is  decidedly  narrow.  This  outer  zone  may  indeed  be 
entirely  missing  since  tissue  comparable  to  it  is  dis- 
tributed elsewhere  in  the  body,  notably  with  ganglia  along 
the  vertebral  column.  The  organ  is  infrequently  the  seat 
of  alterations,  detectable  either  grossly  or  microscopi- 
cally.    Congestion  and  small  hemorrhages  are  rather 


-MEDULLARY  CARCINOMATOUS  PORTION  OF  THYROID.     UNDULATED  GRASS  PARRAKEET 
(MELOPSITTACUS   UNDULATUS). 


THE  DUCTLESS  GLANDS  337 

common  in  acute  infectious  disease  especially  when  the 
respiratory  system  is  involved,  but  these  rarely  destroy 
tissue  or  materially  reduce  the  chromophilic  cells.  These 
circulatory  disturbances  have,  however,  been  predomi- 
natingly among  the  mammals  although  birds  have  suf- 
fered with  infections  to  a  high  percentage.  The  medulla 
is  much  more  often  the  seat  of  congestion  while,  when 
hemorrhage  has  occurred,  the  cortex  is  apparently  always 
involved.  Lipoidal  reduction  has  been  seen  in  a  few 
mammals.  Primates  and  Carnivora,  once  to  a  state  of 
complete  exhaustion.  More  serious  lesions  have  occurred 
eleven  times,  and  since  the  cause  and  meaning  of  disease 
in  this  body  are  so  vague  it  seems  well  to  recite  briefly 
each  one. 

A  Weeper  Cebus  {Cehus  capucinus)  suffered  for 
several  months  with  constantly  but  slowly  increasing 
skeletal  deformity  of  the  osteomalacic  variety.  He  died 
after  moving  him  to  a  new  cage,  his  end  being  hurried  by 
a  scalp  wound.  At  autopsy  the  skeletal  condition  was 
determined  to  be  of  the  above  named  kind.  The  organs 
were  in  good  condition.  The  adrenals  were  decidedly 
enlarged  for  a  monkey  of  this  size,  measuring  1.8  cm.  in 
length.  The  medulla  was  a  solid,  brownish,  homogeneous 
portion  covered  with  a  very  narrow,  barely  discernible 
cortical  zone.  This  was  apparently  due  to  a  uniform 
hypertrophy  of  the  cells  of  the  medulla.  The  testes  were 
slightly  atrophied  and  fibrotic.  A  Black  Spider  Monkey 
{Ateles  ater)  had  a  history  of  stiffness  of  legs  for  six 
months.  This  was  probably  a  sign  of  osteomalacia  since 
at  autopsy  this  condition  was  found  together  with  a 
secondary  anemia,  chronic  gastritis,  acute  enteritis  and 
brown  atrophy  of  heart.  The  adrenal  was  knob-shaped, 
the  cortex  was  wide,  brown,  regular,  the  medulla  small 
gray-purple.  ''Histologically  the  capsule  of  the  adrenal 
is  thicker  than  is  commonly  seen  in  Primates  and  con- 
nective tissue  bands  between  the  units  of  the  zona  glomer- 


338   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

ulosa  are  somewhat  stouter  than  common.  The  cells  of 
this  layer  take  the  stain  a  little  more  deeply  than  usual, 
but  are  otherwise  negative.  The  layer  separating  cortex 
and  medulla  is  occupied  by  a  band  of  well  formed  con- 
nective tissue  which  is  not  proceeding  inward  but 
outward  and  so  encroaching  upon  the  zona  reticularis  as 
to  remove  it  completely  in  places,  in  others  to  make 
isolated  islands  of  its  cell  groups.  Fine  lines  of  connective 
tissue  are  penetrating  from  this  into  the  middle  layer  but 
not  disturbing  it  as  yet.  The  connective  tissue  septa  pene- 
trating the  medulla  are  somewhat  wider  than  one  would 
expect  but  show  no  activity  in  their  growth.  The  medulla 
is  somewhat  broken  up,  vacuolated  and  the  chromophilic 
cells  are  not  especially  prominent,  indeed  some  of  them 
seeming  to  have  undergone  necrosis."  A  puma  {Felis 
concolor)  died  after  a  sickness  of  two  weeks  from  acute 
gastroenteritis  with  its  usual  visceral  associations,  includ- 
ing acute  nephritis,  and  with  calcifications  in  the  adrenals. 
These  structures  were  quite  firm  and  nodular,  on  section 
tough  and  resilient.  ''The  cortex  is  irregular,  brown, 
with  paler  brown  medulla.  Areas  of  calcifications  appear 
as  small  dots,  as  linear  formations  and  in  some  places 
seemingly  around  blood  vessels.  Histological  section 
shows  marked  vacuolization  of  cells,  particularly  of  cor- 
tex. There  is  a  diffuse  overgrowth  of  connective  tissue 
which  has  become  hyaline.  Here  and  there  smaU  cal- 
careous deposits  may  be  seen  but  no  massive  areas  as 
mentioned  above. ' ' 

A  Himalayan  Thar  {Hemitragus  jemlaicus)  came 
to  his  end,  after  a  history  of  convulsions,  from  nephritis, 
which  had  resulted  in  general  edema  including  the 
serous  sacs,  and  an  associated  cardiac  dilatation. 
His  adrenal  was  egg-shape,  of  normal  size,  ^\Tth  a 
wide,  irregular  dull  brown  cortex  and  a  homogeneous 
opaque,  darker  brown  medulla.  ''Histologically  the  cellu- 
lar structure  of  the  cortex  is  partly  destroyed,  partly 


THE  DUCTLESS  GLANDS  339 

dropped  out  and  partly  disturbed  by  overgrowth  of  con- 
nective tissue.  This  connective  tissue  is  quite  prominent 
in  the  medulla  where  it  is  surely  increased  although  it  is 
made  more  prominent  by  absence  of  cells,  some  of  which 
have  been  degenerated  and  some  dislodged. ' '  A  Japanese 
Macaque  {Macacus  fuscatus)  after  drooping  three  weeks 
presented  at  autopsy  the  following  numerous  lesions: 
anemia,  chronic  atrophic  gastritis,  atrophy  of  heart  muscle 
with  regeneration,  hemosiderin  pigmentation  of  liver, 
perilobular  fibrosis  of  liver,  chronic  diffuse  nephritis 
(subcapsular  type),  congestion  of  spleen,  fibrillar  fibrosis 
of  spleen,  hemosiderin  pigmentation  of  spleen,  local 
amyloid  infiltration  of  spleen,  calcareous  infiltration  in 
medulla  of  adrenal.  Grossly  the  adrenal  showed  a  thick, 
orange  yellow  cortex  and  small  solid,  brown  medulla. 
''Histologically  the  organ  appears  noiTual  in  all  respects 
save  for  the  presence  of  a  few  small  irregular  areas  of 
calcification  in  the  medulla.  These  occur  apart  from  any 
recognizable  necrotic  or  fibrous  areas.  In  one  place  one 
appears  to  lie  within  the  lumen  of  a  blood  vessel.  No 
fibroses  or  special  congestions  found  anywhere  in  the 
organ  and  cells  show  normal  details  and  normal  numbers 
of  vacuoles. ' ' 

A  California  hair  seal  (Zalophus  calif ornianus) 
which  had  been  refusing  food  and  having  loose  stools 
for  about  ten  days  presented  after  death  the  fol- 
lowing diagnosis:  Hypernephroma  of  adrenal,  chronic 
hypertrophic  enteritis  with  acute  exacerbation,  hemor- 
rhagic splenitis,  passive  congestion  of  liver,  congestion 
and  edema  of  lungs  with  catarrhal  pneumonia,  acute 
fibrinous  pericarditis,  chronic  lymphadenitis,  chronic 
interstitial  nephritis.  His  right  adrenal  seemed  about 
normal,  being  5x2x1  cm.  with  a  narrow,  dull 
yellow  cortex  and  a  large  mottled  gray  brown  medulla. 
The  left  one  was  5x3x1  cm.  The  upper  pole  is 
swollen   and  contains  in   its  centre   a   spherical   tume- 


340  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

faction  which  is  red,  mottled,  sharply  outlined,  with  a 
suggestion  of  a  capsule  and  slightly  firmer  than  surround- 
ing organ.  ' '  Histological  section  shows  a  capsule  of  very 
noticeable  thickness  but  possibly  not  much  in  excess  of 
normal.  The  cortex  particularly  in  its  deeper  layers  is 
much  injected  and  in  some  places  there  has  been  hemor- 
rhagic diffusion.  In  many  places  in  the  zona  fasciculata, 
more  especially  near  the  periphery,  there  is  breaking  up 
of  the  cell  tubes  with  an  infiltration  of  large  round  cells 
and  some  chromatophilic  cells.  In  other  places  this  seems 
to  have  gone  on  to  fibrous  tissue  increase  and  necrosis  of 
the  cortical  fibrous  cell  types.  The  coim.ective  tissue  layer 
below  the  cortex  is  wide  and  the  spaces  filled  with  blood. 
This  connective  tissue  also  surrounds  islets  of  medullary 
cells  which  are  not  specially  chromatophilic.  The  mass 
in  the  medulla  is  made  up  of  varying  sized  alveoli  sur- 
rounded by  rather  rare,  highly  vascularized  connective 
tissue  and  enclosing  islets  of  medullary  cells.  These 
alveoli  may  be  subdivided  by  septa.  Hemorrhage  has 
occurred  into  many  of  them.  The  individual  cell  masses 
are  made  up  of  groups  of  rather  large  cells  with  illy 
defined  margin,  a  granular,  opaque  but  not  vacuolated 
protoplasm.  They  have  a  bladder-like  nucleus  in  which 
the  centrosome  is  large  and  prominent.  Definite  mitotic 
figures  could  not  be  found,  but  mitosis  is  probably 
present.  A  few  cells  with  double  nuclei  were  seen  and 
one  mth  four.  In  many  of  these  large  islands  the  centre 
has  gone  to  pieces  from  hemorrhage  or  necrosis.  Some  of 
the  vessels  are  thrombotic  and  one  shows  a  ver^^  pro- 
nounced periarteritis. ' '  A  brown  cebus  (Cebiis  fatuellus) 
was  killed  because  of  a  poor  tuberculin  test  chart.  His 
organs  were  negative  except  the  right  adrenal  body  which 
was  3  X  1.5  cm.,  or  four  times  the  size  of  its  fellow.  It 
was  a  tense  body  with  a  smooth,  mottled,  deep  yellow 
surface.  On  section  there  were  deep  yellow  islands 
separated  by  pale  brownish  septa;  the  structure  did  not 


THE  DUCTLESS  GLANDS  341 

resemble  adrenal.  Histological  section  showed  a  hyper- 
nephroma of  vacuolated  cell  type,  roughly  alveolar.  Two 
other  cases,  w^hich  because  of  their  microanatomy  are  to 
be  called  hypernephroma  can  be  added  to  those  just  cited. 
One  occurred  in  an  undulated  grass  parrakeet  {Melopsit- 
taciis  undiilatus) ,  the  other  in  a  black  duck  {Anas 
ohscura) .  As  illustrative  of  this  tumor  the  former  will  be 
cited  in  brief.  ' '  Hypernephroma  of  adrenal  with  hemor- 
rhage into  body  cavity.  A  tumor  approximating  in  size 
the  head  of  the  host  extends  from  the  region  of  the 
internal  genitalia  and  adrenals  lying  more  on  the  right 
than  on  the  left  side  extending  fully  to  cloaca  and  shoving 
all  abdominal  viscera  forward.  It  has  a  pedicle  springing 
from  between  the  two  upper  lobes  of  the  kidney  where 
adrenals  and  internal  genitalia  are  not  distinguishable. 
Tumor  is  coarsely  lobulated,  well  encapsulated,  nowhere 
adherent.  It  has  a  pale,  dirty  yellow  color,  richly  marked 
by  red  lines  of  congested  vessels.  It  is  fairly  soft,  i.e., 
about  consistency  of  normal  liver.  Upon  incising,  the  cut 
surface  bulges  markedly,  is  a  dirty  gray-yellow,  blotched 
with  darker  gray  areas,  shows  no  internal  hemorrhages 
or  markings  of  special  import.  No  metastases  noted  to 
any  other  organ.  Microscopic  section  shows  a  light  cap- 
sule surrounding  the  tissue  of  the  tumor.  The  latter  has  a 
very  scanty  fibrous  reticular  framework  showing  no 
orderly  or  purposeful  arrangement.  Upon  and  between 
the  reticulum,  irregular  and  for  the  most  part  elongated 
collections  of  cells  are  placed.  At  times  these  present  an 
elongated  fascicular  form,  but  this  is  not  by  any  means  a 
prominent  feature.  The  cells  themselves  are  large, 
rounded  or  polygonal,  have  coarsely  granular  cytoplasm 
which  only  in  rare  cases  contains  vacuoles.  Nuclei  of 
these  cells  stain  very  poorly,  but  it  can  be 
made  out  that  they  are  of  large  epithelial  type  and  of 
vacuolar  appearance. ' ' 

A  somewhat  unusual  tumor  was  encountered  in  a 
Polar  bear  {Ursus  maritimus),  an  adenocarcinoma  of  the 


342   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

adrenal,  when  judged  purely  by  its  histology  but  a  second- 
ary tumor  in  the  lung  displayed  the  more  familiar  picture 
of  large  vacuolated  cells  as  seen  in  hypernephroma.  The 
diagnosis  follows:  *' Scirrhous  adenocarcinoma  of 
adrenals,  secondary  carcinoma  of  lymph  glands,  second- 
ary hypernephroma  of  lung,  secondary  carcinoma  of 
diaphragm,  acute  mucopurulent  bronchitis,  acute  catarrhal 
enteritis,  chronic  diffuse  fibrous  cholecystitis,  choleli- 
thiasis, slight  acute  interstitial  pancreatitis,  follicular 
hyperplasia  of  spleen  with  fibrosis,  hydrothorax, 
hydropericardium,  chronic  hypertrophic  osteoperiostitis, 
encysted  trichina  in  diaphragm,  fatty  infiltration  of 
diaphragm,  chronic  diffuse  nephritis,  chronic  productive 
lymphadenitis,  pigmentation  of  lymph  gland.  Both 
adrenals  are  smaller  than  normal,  of  woodeny  consist- 
ency, the  pale  cortex  and  medulla  are  poorly  separated 
from  each  other.  The  cut  surface  shows  gray  white  and 
tawny  mottling  and  occasional  calcareous  points.  Histo- 
logical section  shows  an  extra  capsule  of  fibrous  tissue 
containing  highly  distended  veins;  the  lining  contains 
masses  of  tumor  cells,  many  of  which  are  necrotic.  It  is 
distinctly  denser  in  type  than  normal  and  more  abundant 
in  places  showing  a  proceeding  fibrosis.  Parenchyma 
shows  but  few  irregular  islands  containing  non-neoplastic 
cells,  some  of  which  are  highly  vacuolated,  others  are  not. 
Interstitial  tissues  in  peripheral  parts  are  often  grown 
together  with  the  deeper  portion  where  are  intermixed 
tumor  areas.  The  latter  consist  of  small  round  acini  of 
variable  size  lined  by  cells  of  active  type.  Nuclei  are 
large  and  hyperchromatic,  cytoplasm  broad  and  disinte- 
grating. Parts  show  necrosis  and  hemorrhage.  Upon 
search  transitionals  from  non-neoplastic  to  neoplastic 
cells  can  be  discovered  in  same  fasciculus." 


SECTION  XII 
THE  SKELETON  AND  ITS  JOINTS 

The  bones  with  their  articulations  have  been  the 
subject  of  extensive  study  and  research  by  zoologists  in 
the  direction  of  classification  and  evolution.  Adaptation 
of  the  osseous  construction  to  the  needs  of  the  animal  is 
well  appreciated  biological  knowledge.  For  example,  the 
keel  of  the  sternum  in  birds  affords  broad  origins  for  the 
flying  muscles,  the  pectorals,  which  also  insert  on  the 
ala3  of  this  bone  and  on  the  clavicle,  and  in  addition  use 
these  latter  formations  as  fulcra.  So  too  the  extremities 
of  quadrupeds  are  angular  in  their  upper  two  segments 
for  the  purpose  of  supplying  a  direct  action  of  the  flexors 
employed  in  running  and  leaping.  The  thick  masseter 
muscle  of  carnivores  is  accommodated  in  the  deep  zygo- 
matic fossa.  Many  other  examples  might  be  cited,  but 
these  serve  to  direct  attention  to  the  adaptation  of 
function  and  construction.  Pathological  changes  in 
our  materials  are  however  too  few  to  permit  con- 
clusions as  to  possible  relation  of  zoological  position 
and  development  except  such  as  may  refer  to  deformity 
incident  to  the  degenerative  processes — rickets,  osteo- 
malacia and  osteogenesis  imperfecta,  and  in  these 
conditions  the  alterations  are  merely  passive  accommoda- 
tions to  weakened  support  in  order  to  obtain  comfort.  To 
put  the  matter  in  other  words,  it  would  seem  that,  aside 
from  the  diseases  just  named,  there  is  no  outstanding 
change  in  the  skeletal  tissues  peculiar  to  zoological  orders 
that  might  indicate  vulnerability  of  the  system  or  the 
methods  of  response  to  injury  or  disease. 

Effects  of  Trauma. 

There  must  be  considerable  reserve  or  reconstructive 
power  in  the  bones  of  animals  since  it  is  a  common  thing 
at  autopsy  to  see  unmistakable  evidences  of  repair  of 

343 


344  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

fractures,  dislocations  and  inflammations.  Some  illus- 
trations are  introduced  to  exemplify  this  healing  ability, 
one  of  which  was  found  in  an  animal  shot  by  a  hunter, 
the  other  an  incidental  autopsy  discovery.  Even  though 
there  be  no  definite  relationship  between  the  zoological 
order  and  osseous  disease,  it  is  interesting  to  record  a 
very  simple  observation.  Animals  with  long  extremities, 
especially  when  the  bones  are  quite  near  the  skin,  have  a 
rather  high  incidence  of  fractures  and  inflammations. 
Thus  the  ungulates  have  of  all  orders  the  highest  per- 
centage of  these  traumatic  and  infective  lesions ;  herons 
and  gallinaceous  birds  follow  the  ungulates.  Marsupials, 
primates  and  carnivores,  in  this  order,  are  susceptible  to 
inflammations  but  not  to  fractures.  Bones  are  often 
broken,  among  the  Cervidse,  Bovidse  and  Camelidae,  when 
as  they  are  chased  by  mates,  they  fall  upon  the  slippery 
floor  of  the  cages;  or  again  the  mounting  of  a  small 
animal  by  a  large  buck  may  crush  the  former  to  the  earth. 
Two  cases  of  fractured  pelvis  have  been  seen  in  antelopes 
from  a  fall  with  extended  hind  legs. 

It  would  seem  that  repair  is  usually  satisfactory  if  the 
animal  have  a  quiet  retreat  where  callus  may  form  and 
union  occur.  A  heron  is  known  to  have  broken  both  bones 
of  the  leg ;  at  autopsy  a  very  insignificant  circumferential 
callus  remained,  the  member  being  as  straight  and  strong 
as  normal.  Figure  40  shows  the  femur  of  a  deer  shot  by 
a  hunter ;  the  shortening  was  considerable,  but  function 
was  doubtless  good  because  the  hunter  could  perceive  no 
limping  as  the  animal  ran.  The  most  interesting  fracture 
among  our  records  was  an  intracapsular  fracture  of  the 
hip  in  a  Huanaco  (Lama  huanacos)  shown  in  Figure  41. 
This  animal  slipped  on  the  ice  in  December  and  was 
thought  to  have  broken  something  near  the  hip,  but  it 
limped  around  mthout  any  great  show  of  pain  until  the 
following  May,  when  it  died  of  meningitis  secondary  to  an 
otitis  media  et  interna.  At  autopsy  an  unhealed,  complete 
fracture   of  the   neck   of  the   right   femur   was   found. 


Fic;.  40. — HEALED   FRACTURE  OF  FEMUR.     FROM  A  DEER  SHOT  BY  A  HUNTER. 


Fig.   41.— partially   HEALED   INTRACAP.SLLAR   KRAC  TL  RK   OE   HEAD   OF 
RIGHT   FKMIR.      HLANACO    <I.A\L\    HIANACOSI. 


THE  SKELETON  AND  ITS  JOINTS  345 

apparently  separating  the  head  from  the  neck,  the  former 
being  dislocated  to  the  upper  angle  of  the  obturator  fora- 
men. Everywhere  about  the  joint  callus  had  been  thrown 
out,  but  not  in  a  manner  to  effect  a  junction  of  the  broken 
ends  nor  to  seal  the  edge  of  the  acetabulum  to  the  femoral 
neck.  This  was  probably  due  in  part  to  the  irregularity 
of  the  line  of  fracture  and  to  the  interposition  of  the 
upper  part  of  the  dislocated  head  between  the  lower  rim 
of  the  acetabulum  and  the  surgical  neck  of  the  bone. 
"When  the  specimen  was  fresh  traces  of  capsule  were 
found  over  the  upper  half  of  the  acetabulum.  While  it  is 
usually  difficult  to  decide  the  manner  in  which  these 
injuries  effect  their  damage  and  deformity,  it  might  be 
ventured  to  explain  this  case  as  due  to  extreme  postero- 
lateral extension  of  the  leg  driving  the  head  of  the  femur 
downward  and  inward,  rupturing  the  capsule  and  the 
ligament  bridging  the  acetabular  notch,  to  rest  on  the 
pubis  at  the  upper  edge  of  the  obturator  foramen  where 
it  could  find  a  sort  of  joint  cavity  made  by  the  pubic  and 
ischial  segments  of  the  old  acetabulum,  but  about  an  inch 
and  a  half  below  its  normal  location. 

Another  injury  to  the  hip  joint  was  noted  in  a  Living- 
stone's eland  {Taurotragus  oryx  livingstonii).  This 
beast  was  not  positively  known  to  have  fallen,  although  it 
was  suspected  that  such  an  accident  had  occurred  by 
reason  of  sudden  inability  to  rise.  At  autopsy,  death 
having  succeeded  on  signs  of  shock,  a  complete  upward 
and  backward  dislocation  of  the  right  femoral  head  was 
found ;  there  was  also  an  intracapsular  rupture  of  the  left 
round  ligament,  but  on  this  side  the  femoral  head  had  not 
left  the  acetabular  cavity. 

Many  other  fractures  have  been  observed  but  gen- 
erally mthout  interesting  features.  The  conclusions 
which  may  be  drawn  from  our  experience  are  that 
animals  with  long  bones,  and  liable  to  chase  have  the 
greatest  liability  to  fractures,  and  that  the  healthy  beast, 
23 


346   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

given  seclusion  and  quietude,  possesses  great  ability  to 
heal  its  broken  bones.  Pathological  fractures  are 
occasionally  seen.    (Consult  notes  on  cretin  wolves.) 

Before  entering  upon  a  discussion  of  the  most  impor- 
tant of  osseous  lesions,  rickets  and  osteomalacia,  certain 
inflammatory  states  may  be  appropriately  described. 

Inflammations. 

H37)ertrophic  osteoperiosteitis :  A  male  lion  (Felis 
leo)  at  the  Garden  three  years  died,  after  being  out  of 
condition  for  a  long  time,  from  chronic  ulcerative  pul- 
monary tuberculosis  \vith  terminal  pneumonia,  nephritis 
and  enteritis.  Both  hind  feet  had  been  observed  as 
enlarged  and  apparently  painful  for  some  weeks  before 
death.  Upon  dissection  the  bones  of  both  hind  feet  are 
the  seat  of  extensive  hypertrophy,  and  the  periosseous 
fibrous  tissues  are  thickened.  A  large  mass  about  the  size 
of  a  small  orange  lies  attached  to  the  outer  side  of  each 
ankle.  The  hypertrophic  periosteitis  extends  up  the 
tibia  a  distance  of  about  three  inches  and  the  fibula  for 
about  the  same  distance.  These  two  bones  are  adherent 
to  each  other  for  about  li/o  inches.  The  joint  between 
them  and  the  tarsal  bones  is  apparently''  perfectly  free. 
The  calcaneum  is  the  bone  most  severely  involved ;  on  this 
is  a  large  rounded  mass  which  extends  on  the  bone  for  a 
distance  of  about  21/0  inches.  The  small  bones  of  the  foot 
are  more  or  less  severely  involved  but  are  not  bound 
together,  the  joints  being  practically  free.  The  terminal 
and  next  phalanges  are  entirely  free  from  disease  while 
the  metatarsals  are  severely  involved  and  grown  together 
into  one  large  mass.  On  section  this  appears  as  a  mass  of 
spongy  bone  lying  on  top  of  the  cortex.  In  the  dried 
specimen  this  looks  very  like  old  pumice  stone.  Histo- 
logical section  shows  the  periosteum  raised  from  the  bone 
by  mononuclear  infiltration.  The  bone  marrow  spaces  are 
filled  by  a  very  delicate  gelatinous  material.  The  lameUas 
are  thickened.    A  photograph  of  the  foot  with  a  normal 


Fii;.    42.— HYPHRTROPHIC     PKRIOSTEITIS.       RIGHT     HIND     FOOT    WITH    A    NORMAL    LEI  1.      1.1( 
(FELIS  LEO).    THIS   CONDITION   WAS  ASSOCIATED    WITH   CHRONIC   PULMONARY  TCBERCL  LOS 


Fir,.  43. — MARKED  SCOLIOSIS  IN   A  COCKATOO. 


THE  SKELETON  AND  ITS  JOINTS  347 

example  is  given.  (Fig.  42.)  (See  also  Tuberculosis 
section — Garni  vora. ) 

A  cockatoo  died  from  acute  miliary  tuberculosis ;  the 
upper  thoracic  and  lower  cervical  vertebras  are  involved 
in  an  S-shaped  scoliosis  which  reduces  the  height  of  the 
thorax  by  perhaps  a  centimetre.  Thorough  dissection 
was  not  made,  the  trunk  being  kept  as  a  museum  specimen 
and  for  study  in  event  another  avian  scoliosis  occurred ; 
but  from  palpation,  separation  of  the  muscles  and 
stretching  of  the  spinal  column  it  does  not  appear  that  a 
tuberculous  osteitis  of  the  vertebra  existed.  It  seems 
that  this  may  be  due  to  congenital  deformity  or  old  injury. ' 

A  white-nosed  coati  (Nasiia  narica)  suffered  \m.i\\  gen- 
eralized tuberculosis  which  also  affected  the  wrist  joint 
with  a  caseous  and  ulcerative  arthritis. 

Gouty  arthritis  has  been  recorded  but  three  times, 
although  on  several  occasions  small  uratic  deposits  in 
tendon  sheaths  have  been  observed  in  birds ;  gout  has  not 
been  seen  in  mammals.  An  illustrative  case  in  a  Boat- 
billed  Heron  {Cancroma  coclilearia)  will  be  given  in  the 
section  on  gout. 

Arthritis  as  an  acute  infectious  disease  such  as 
rheumatism  of  the  human  being,  has  not  been  observed, 
but  copious  examples  of  acute,  subacute  or  chronic  mono- 
articular inflammation  are  recorded.  Nearly  all  of  these 
have  a  definite  explanation — traumatism  or  acute  general 
disease,  and  there  are  a  few  cases  of  polyarthritis  with 
chronic  disease.  Notable  among  the  last  are  two  instances 
of  chronic  dry  ossifying  arthritis  and  synovitis,  one  with 
tuberculosis,  the  other  with  actinomycosis,  both  occurring 
in  ungulates.  A  third  case  similar  in  character  deserves 
special  mention.  The  Indian  elephant  ''Bolivar" 
{Elephas  indicus)  died  from  pulmonary  tuberculosis, 
myocarditis,  nephritis  and  hepatic  cirrhosis.  The  joints 
of  all  extremities  showed  atropliic  arthritis  with  fluid,  the 
synovial  membranes  being  ulcerated  or  retracted  and 
fibrotic.    The  articulating  surfaces  wiiere  not  roughened 


348   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

by  erosion,  were  flattened.  It  is  perhaps  worthy  of 
mention  that  this  old  and  familiar  animal  was  the  occu- 
pant of  the  same  enclosure,  floored  with  cement,  for  over 
thirty  years,  conditions  which  might  be  partly  instru- 
mental in  the  arthritic  changes  as  well  as  in  the  flattening 
of  articular  surfaces. 

The  Ungulata  frequently  suffer  with  wounds,  ulcers 
and  abscesses  about  the  lips,  nose,  and  soft  tissues  of  the 
jaws  which  may  at  times  be  confusingly  like  actinomy- 
cosis. This  disease  we  have  seen  in  gazelles  and  tapirs 
but  have  had  to  exclude  it  in  several  other  members  of  this 
order.  A  number  have  come  to  autopsy  with  osteitis  of 
the  lower  mandible,  some  evidently  traumatic  in  origin, 
others  probably  due  to  infection  via  the  teeth.  Figure  44 
represents  the  jaw  bone  of  an  Isabelline  gazelle  {Gazella 
Isabella)  suffering  with  a  rarefying  osteitis  from  a  root 
abscess,  and  illustrates  well  the  possibility  of  focal  infec- 
tion from  this  source. 

Degenerative  Skeletal,  Diseases. 

While  the  foregoing  instances  of  disease  in  the  osseous 
system  are  interesting  examples  of  individual  patho- 
logical lesions,  they  are  insignificant  in  comparison  with 
the  forms  of  bony  change  known  under  the  names  of  rick- 
ets, osteomalacia,  osteogenesis  imperfecta  and  the  like- 
systemic  conditions  which  are  chiefly  degenerative  but 
have  certain  evidences  of  inflammation  in  addition.  The 
modern  knowledge  of  the  first  two  named  is  so  far  from 
complete  that  it  cannot  be  said  that  there  is  any  certainty 
of  their  identity.  Indeed  there  seem  to  be  some  reasons  to 
think  that  there  is  more  than  one  variety  of  rickets,  that  all 
cases  are  not  dependent  upon  the  same  cause,  and  that  in 
essence  it  is  the  same  process  as  osteomalacia,  the  latter, 
however,  occurring  at  a  later  age.  We  shall  show  that  in 
the  same  order,  Primates,  both  diseases  may  occur  in 
animals  fed  upon  the  same  diet,  and  that  one  family  tends 
to  have  one  disease,  another  family  the  other. 


Fig.   44. — DENTAL  ROOT  ABSCESS  AND   OSTEITIS  OF  JAW   BONE. 
ISABELLINE  GAZELLE   (GAZELLA  ISABELLA). 


THE  SKELETON  AND  ITS  JOINTS  349 

ElCKETS. 

Since  the  two  conditions  are  diagnosed  separately  in 
veterinary  practice  and  each  seems  to  have  a  distinct 
place  in  medical  ideas,  it  may  be  well  to  outline  upon  what 
criteria  the  two  diagnoses  have  been  made  in  this  Garden. 
Rickets  is  essentially  a  disease  of  early  life.  The  animal 
is  noted  as  having  a  large  head,  squatty  station,  heavy 
extremities  and  a  prominent  belly.  Death  occurs  as  the 
result  of  enteritis  or  pneumonia.  Occasionally  such  a 
young  specimen  seems  to  recover  from  the  disease  but 
retains  the  distortion  of  his  skeleton;  this  is  important, 
for  we  believe  that  osteomalacia,  except  the  variety  con- 
iined  to  periods  of  pregnancy,  rarely  ends  in  recovery 
when  once  thoroughly  established.  At  autopsy  the  cranial 
bones  are  the  seat  of  osteotabes,  the  face  is  broad,  the 
epiphyseal  junctions  are  swollen  by  irregular  osteo- 
genesis and  granulation  tissue,  the  periosteum  shows 
an  irregular  fibrous  tissue  overgrowth — the  last  two 
processes  producing  bones  of  irregular  contour  and 
thickness.  Section  through  the  osteogenetic  ends  of  the 
long  bones  shows  actively  congested  marrow  up  to  the 
articular  cartilages  with  very  tortuous  strands  of  spongy 
bone  or  cartilage,  and  when  considered  transversely, 
there  is  a  bone-forming  layer  of  many  times  the  normal 
thickness  but  bloody  red  instead  of  pink. 

Osteomalacia. 

Osteomalacia  appears  in  mature  animals  or  at  least 
those  well  able  to  care  for  their  own  nourishment.  The 
earliest  observations  are  not  referable  to  the  skeleton  but 
to  the  change  in  the  activity  of  the  beast.  He  will  be  noted 
as  less  active  in  running,  jumping  or  searching  for  his 
food.  The  customary  position  is  a  sitting  or  lying  one. 
No  change  is  noted  in  the  head  or  face.  As  the  disease 
progresses,  the  animal  becomes  quite  inactive,  seeks  soli- 
tude but  will  eat  well  if  the  food  be  conveniently  available 
and  he  does  not  have  to  fight  for  it.    The  movements  are 


350   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

stiff  and  seem  painful.  About  this  time  definite  alteration 
in  the  shape  of  the  chest  is  perceptible,  and  in  some  cases 
there  is  anterior  curvature  of  the  legs.  Movement 
becomes  so  difl&cult,  probably  from  weakness  and  pain, 
that  it  seems  as  if  paraplegia  actually  existed.  The 
inability  of  affected  monkeys  to  climb  has  given  rise  to  the 
term  ''cage  paralysis,"  but  this  term  should  not  be 
restricted  to  weakness,  the  result  of  osteomalacia  since  it 
is  used  by  dealers  and  keepers  to  imply  the  cramped 
station  and  gait  of  an  animal  long  housed  in  quarters  too 
small  for  it,  an  appropriate  application  because  it  sug- 
gests cause  and  effect.  However,  the  appellation  is  \\ddely 
and  loosely  used  insuring  its  employment  in  diagnosis 
for  entirely  different  conditions  such  as  degenerative 
bone  disease  and  hind-quarter  laming  from  enteric  intoxi- 
cations ;  for  these  affections  one  might  use  the  term  in  an 
adjectival  or  descriptive  sense. 

Our  Primate  collection  has  suffered  considerably  with 
osteomalacia,  and  we  have  devoted  much  time  to  the  study 
of  its  cause  and  treatment.  However,  the  Garden  is  not 
alone  in  this  experience,  for  wherever  certain  species  are 
kept  the  disease  appears.  The  description  of  cases  in  the 
New  York  Garden  by  Blair  and  Brooks  (1)  is  excellent,  and 
with  the  exception  of  data  concerning  the  nervous  system, 
almost  exactly  parallels  our  own  observations.  They  lay 
much  stress  upon  the  changes  in  the  brain,  cord  and 
ganglia  as  constant  in  well  developed  cases  but  as  prob- 
ably secondary  to  the  osseous,  hemic  and  metabolic 
disturbances.  We  have  been  unable  to  find  any  patho- 
logical lesions  in  four  thoroughly  studied  brains  and 
cords  from  well  developed  cases.  As  will  appear  later, 
our  most  satisfactory  findings  were  in  the  dietaiy  and 
metabolic  chemistry  and  in  the  osseous  pathology.  The 
cases  recorded  by  Campbell  and  Cleland(2)  would  seem 

( 1 )  See  Blair  and  Brooks,  Osteomalacia  of  Primates  in  Captivity, 
Ninth  Annual  Report,  New  York  Zoological  Society,  1904,  p.   135. 

( 2 )  Campbell  and  Cleland,  Jour.  Comp.  Path,  and  Ther.,  Vol.  32,  p.  95. 


THE  SKELETON  AND  ITS  JOINTS  351 

to  be  undoubted  instances  of  myelitis,  but  the  osseous 
changes  are  not  sufficiently  discussed.  In  many  cases  it 
would  seem,  therefore,  that  there  is  some  change  in  the 
nervous  system,  but  there  may  be  some  examples  without 
this  and  with  predominant  osseous  lesions.  We  are  in- 
clined to  think  that  these  two  groups  differ  quahtatively, 
and  we  look  upon  the  confusion  as  demanding  for  its  ulti- 
mate solution  the  use  of  exact  nomenclature,  especially  the 
exclusion  of  "cage  paralysis"  as  a  diagnostic  term.  The 
only  division  we  can  understand  at  the  present  time 
depends  upon  the  gross  changes  in  the  bones,  those  with 
and  those  without  definite  irregularities  in  contour  due 
to  periosteal  overgrowth.  Certain  of  the  former  may 
show  no  unevennesses  at  all,  the  deformity  being  due  to 
softness  of  the  skeleton.  The  other  group  has  shafts  of 
irregular  thickness,  swellings  around  the  joints  and  much 
beading  of  the  middle  of  the  ribs. 

Whether  or  not  there  be  true  paralysis  is  difficult  to 
settle,  but  in  our  cases  we  have  decided  always  in  the 
negative  because  of  the  ability  of  the  monkeys  to  grasp 
firmly  with  the  hind  digits.  The  animals  tend  to  lie  in  one 
position,  determined  probably  by  comfort,  the  result 
being  that  they  develop  sores  at  the  points  of  contact  with 
their  cage  floor,  and  deformities  of  the  skeleton  (see 
Fig.  45).  These  deformities  are  especially  well  exhibited 
by  the  chest,  the  vertebrae  and  the  pelvis  and  are  referable 
to  the  almost  constant  squatting  of  the  animal ;  the  long 
bones  may  be  bowed  but  not  as  much  as  in  rickets,  nor  is 
the  epiphyseal  junction  so  knobby  as  in  that  disease. 

The  foregoing  description  is  based  chiefly  upon  obser- 
vations on  monkeys  but  may  be  closely  paralleled  in 
carnivores  and  rodents.  These  latter,  however,  lie  rather 
than  sit  during  the  development  of  the  disease,  so  that 
thoracic  and  pelvic  deformity  is  relatively  less  than  in 
monkeys.  Death  is  due  to  enteritis,  anemia,  shock  from 
fractures  and  respiratory  inflammation. 


352   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

In  so  far  as  the  deformities  of  the  skeleton  may  serve 
to  distinguish  between  rickets  and  osteomalacia,  I  can 
only  point  to  the  preponderance  of  changes  in  the  skull 
and  extremities  in  the  former  and  of  the  trunk  bones  in 
the  latter.  Deformity  of  the  chest,  barrel-shape  short- 
ening and  pigeon  breast,  is  due  more  to  posture  than  to 
the  essentially  osseous  changes.  ''Rachitic  rosary"  may 
occur  in  both,  but  it  is  always  better  exhibited  in  rickets ; 
in  this  disease  the  swellings  occur  at  the  costochondral 
junction,  while  in  osteomalacia  rosary-like  nodules  may 
develop  anywhere  along  the  ribs. 

Examination  of  the  anatomical  lesions  is,  however, 
somewhat  more  helpful,  and  the  following  description  for 
osteomalacia  may  be  contrasted  mth  that  already  given 
for  rickets.  The  peculiar  change  is  a  thinning  of  the  shaft 
of  long  bones  and  reduction  of  the  subperiosteal  plates 
of  flat  bones. 

In  mammals  the  long  bones  are  more  affected  than  in 
birds  whose  sternum,  ribs  and  beak  show  the  severest 
changes.  The  skull  is  frequently  not  affected  to  a  serious 
degree,  but  may,  however,  show  advanced  lesions,  the 
cranial  plates  being  thinned  in  places  so  that  they  may 
be  bent  in,  or  occasionally  a  periosteal  thickening  may 
be  found ;  the  head  as  a  whole  is  not  misshapen.  The  ribs 
are  softened  and  may  be  of  paper  thickness  although 
there  may  be  found  a  periosteal  overgrowth,  perhaps  a 
kind  of  splinting,  which  makes  the  diameter  variable.  At 
costochondral  junctions,  beading  may  be  found,  but  with- 
out the  active  congestion  seen  in  rickets.  Similar 
alterations  may  be  found  in  the  long  bones,  here  in 
characteristic  degree  in  that  the  shaft  walls  are  thin,  by 
removal  of  the  endosteal  and  periosteal  layers  sometimes 
Avith  definite  retraction  of  the  marrow.  Occasionally  sub- 
periosteal thickenings,  made  of  osteofibrous  tissue  are 
encountered.  At  the  epiphyses  there  are  strands  of 
gelatinous  tissue,  fibrous  and  cartilaginous,  separating 


Fig.   45.— osteomalacia.     MODERATELY  ADVANCED    CASE   WITH    HOWEVFR  WELL  ESTAB- 
LISHED    DEFORMITY    OF    THORAX    AND     PELVIS.      THIS     POSITION    WAS    CONSTANT    FOR    THREE 
BLACK   HANDED  SPIDER  MONKEY   (ATELES  GEOFFROYI). 


MONTHS   BEFORE  DEATH. 


THE  SKELETON  AND  ITS  JOINTS  353 

piiik  or  blood  red  areas  of  marrow.  These  strands  may 
contain  calcareous  matter  and  are  probably  the  remains  of 
the  cancellated  tissue.  Despite  all  this  activity  at  the 
ends  of  the  long  bones  there  is  not  the  extreme  promi- 
nence of  articulations  so  characteristic  of  rickets. 
Gelatinous  or  cartilaginous  islands  may  be  seen  in  the 
deep  red  shaft  marrow. 

Fractures  may  be  found  and  around  them  may  form  a 
blood  clot  or  loose  fibrous  tissue  entirely  devoid  of  bone 
salts.  If  a  break  has  existed  for  some  time  a  very  pro- 
nounced fibrous  overgrowth  from  the  periosteum  is  apt 
to  occur,  indeed  an  excessive  fibrosis  may  exist,  but  this  is 
ineffective  for  healing  of  the  fracture  or  splinting  of  the 
shaft.  Certain  cases,  notably  in  Carnivora,  seem  to  have 
especial  activity  in  and  around  joints  so  that  when  the 
member  is  dissected  one  gets  the  impression  of  osteoar- 
thritis. In  such  cases  the  synovia  may  be  fibrotic  and  the 
articular  surfaces  dry./  The  pelvic  deformities  are  simi- 
lar to  those  in  the  hmnan  being — lateral  contraction  with 
bending  in  of  the  superior  rami  of  the  pubis  with  the  pro- 
duction of  a  beak,  to  which  the  name  ^'duckbill"  has 
been  given.  The  anterior  curvature  of  the  lumbar  spine 
makes  an  acute  angle  at  the  upper  end  of  the  sacrum. 

Histological  examination  of  a  number  of  our  cases  of 
osteomalacia  and  rickets  have  failed  to  show  any  lesion 
different  from  those  known  for  the  human  being  and  for 
domesticated  animals.  It  is  noteworthy  that  not  all  bones 
of  a  given  case  will  show  the  changes  to  the  same  degree 
even  though  grossly  they  may  seem  comparably  affected. 
So  too  there  is  no  certain  relation  between  the  degree  of 
deformity  as  shown  by  the  body  as  a  whole  and  the 
advancement  of  osteoporosis  as  seen  under  the  micro- 
scope. These  observations  are  in  accord  with  those  of 
Brooks  and  Blair.  Just  why  this  is  cannot  be  stated,  but 
as  the  cause  of  these  two  bone  diseases  may  not  always 
be  the  same,  variations  in  gross  and  minute  anatomy  are 
not  remarkable. 


354   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

t  Analysis  of  the  bones  shows  a  loss  of  calcium  and  an 
excess  of  sulphur  and  magnesium. "^  The  loss  of  the  first 
is  chiefly  via  the  intestinal  discharges  but  also  via  the 
urine.  The  metabolism  of  one  monkey  showed  a  liigh 
calcium  and  phosphorus  loss  with  moderate  retention  of 
sulphur  and  magnesium. 

Because  of  the  importance  of  osteomalacia  and 
rickets  in  cebus  monkeys  and  certain  other  animals,  Dr. 
E.  P.  Corson-White  has  been  investigating  its  etiology. 
I  shall  refer  briefly  to  her  results  as  they  affect  our  pres- 
ent subject  but  shall  leave  for  discussion  in  the  chapter 
on  diet,  which  she  has  written,  the  broader  question  of 
food  and  systemic  disease. 

It  must  be  understood  that  the  instances  included  in 
this  general  discussion  of  degenerative  osseous  disease 
are  cases  of  definite  character  and  development.  There 
may  have  been,  in  addition  to  the  numbers  cited  in  the 
list  on  page  357,  many  more  animals  at  autopsy  with 
early  or  unrecognized  constructive  or  destructive  abnor- 
malities, and  we  are  thoroughly  familiar  with  the 
imperfect  skeletal  development  of  specimens,  inbred  or 
reared  in  captivity  or  even  those  adult  when  caught  yet 
under  Park  conditions  for  many  years.  In  these  latter 
groups  the  changes  vary  from  incomplete  construction 
(an  example  of  atrophy  was  quoted  on  page  24)  to  actual 
degeneration  as  in  osteomalacia.  Inbreeding  seems  to  be 
a  potent  factor  in  many  cases,  a  well  known  fact  in  hmnan 
and  veterinary  medicine.  The  importance  of  inactivity 
in  the  causation  of  degenerative  bone  disease,  the  unused 
muscles  giving  the  bones  nothing  to  do,  is  certainly  admit- 
ted but  it  is  immeasurable.  It  is  probably  not  great  in  a 
cage  of  mixed  varieties  of  monkeys.  The  effect  of  the 
absence  of  sunlight  in  osseous  degeneration  is  no  factor 
in  our  material.  The  exhibition  house  is  well  lighted  and 
many  animals  are  out  of  doors  all  year  around. 

The  ductless  glands  have  repeatedly  been  accused  of 
responsibility  for  these  disorders.    In  our  seventy-nine 


THE  SKELETON  AND  ITS  JOINTS  355 

cases  of  osteomalacia  and  thirty-four  of  rickets,  no 
abnormality  has  been  observed  in  ovary,  testes  or 
adrenals  except  for  moderate  congestions.  Two  cases 
were  associated  with  pancreatic  disease,  once  acute,  once 
chronic.  The  thyroid  body  has  been  found  to  have  been 
definitely  abnormal  only  once — secondary  hyperplasia 
with  colloid  in  a  carnivore.  In  the  Primates  this  body  was 
frequently  congested  and  has  shown  small  colloid  cysts 
but  was  not  uniformly  enlarged  or  atrophic.  As  a  thera- 
peutic measure  I  have  administered  adrenalin  to  two 
monkeys,  one  for  a  few  weeks,  one  for  nearly  six  months ; 
this  treatment  was  without  any  perceptible  effect  upon 
the  process. 

Dr.  Corson- White  has,  by  the  study  of  some  cases 
during  the  life  of  the  monkey,  confirmed  the  decreased 
alkalinity  of  the  blood  in  connection  with  the  increased 
output  of  calcium  in  the  feces  and  urine. 

Since  the  explanation  of  the  disease  by  blaming  the 
ductless  glands  has  failed.  Dr.  Corson- White  has  under- 
taken a  study  of  the  diet  given  our  monkeys  to  see  if  any 
fault  in  it  were  a  part  of  the  etiology.  Analysis  of  this 
diet  (see  list  page  426)  computed  from  Atwater's  table, 
and  by  actual  analysis  of  the  amounts  of  food  consumed 
by  the  animal  in  four  four-day  periods,  gave : 

1.  Protein— low  in  quantity  and  poor  in  quality ;  espe- 
cially low  in  phosphorus  content. 

2.  Fat — ^very  low. 

3.  Carbohydrate— very  high,  almost  eleven  times  the 
value  of  all  other  ingredients. 

4.  Ash— decidedly  low  and  predominatingly  acid. 
Further  analysis  of  this  ash  showed  a  trace  only  of  cal- 
cium and  phosphorus  and  iron  and  only  a  small  amount 
of  sodium;  potassium,  sulphur  and  magnesium  were 
slightly  higher. 

5.  Vitamines  A,  B,  C,  were  present  in  extremely  small 
amounts— A  was  exceptionally  deficient,  and  in  the 
rations  of  some  days  was  entirely  lacking. 


356  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

There  are  in  this  monkey  diet  several  factors  of 
importance.  1.  Low  vitamine  contents — especially  Vita- 
mine  A — factors  which  are  essential  for  life  and  growth. 
2.  A  high  carbohydrate  diet — which  in  oxidation  yields 
an  acid  ash  and  which  favors  the  growth  of  intestinal 
bacteria  producing  acid  and  gas.  The  acid  from  these  two 
sources  must  be  neutralized  either  by  the  alkali  derived 
from  food,  or  from  the  body  storage.  This  diet,  however, 
is  abnormally  low  in  ash  and  especially  in  the  alkaline 
salts  of  the  ash,  therefore  making  it  an  ideal  diet  for  the 
production  of  osteomalacia. 

The  following  table  shows  the  additions  necessary  for 
corrections  of  the  separate  ingredients  of  the  diet : 


Phos- 


Monkey  diet 

Corrected  by 

Kiee 

Casein,  Salt  mixture,  Carrots,  or  Lettuce. 

Bread 

Casein,   Butter   fat.    Salt  mixture,   especially 

phorus. 

Potato 

Salt  mixture,  especially  Na  CI  and  Ca  C03. 

Raw  peanuts 

Salt  mixture. 

Bananas 

Casein,  Yeast,  or  Carrots. 

Com 

Casein,  Ti-yptophan,  Lacto-albumin. 

Apple 

Casein,  Gelatin,  Butter  fat. 

Onion 

Casein,  Gelatin,  Butter  fat. 

The  complete  diet  may  therefore  be  rendered  adequate  by  the 
addition  of  fresh,  whole  milk  and  leafy  vegetables,  or  by  ))utter  fat, 
salt  mixtures  and  leafy  vegetables. 

It  would  seem  from  these  data  that  in  this  inefficient 
diet  we  have,  if  not  the  cause  of  osteomalacia,  at  least  a 
very  potent  factor  in  its  production.  The  disturbance 
of  the  calcium  and  phosphorus  metabolism  may  be  due 
primarily  to  the  deprivation  of  the  alkaline  salts  from 
the  diet  (famine  osteomalacia)  or  to  a  drain  from  the 
alkaline  storage  of  the  body,  associated  with  a  deficient 
diet  (as  in  the  cases  of  osteomalacia  of  pregnancy  and  lac- 
tation) or  in  the  combined  action  of  a  diet  faulty  in  more 
than  its  salt  content,  which  by  the  production  of  acid  in 
its  oxidation  and  by  favoring  the  development  of  acid- 
forming  bacteria,  causes  the  drain  of  the  body  alkali  for 


THE  SKELETON  AND  ITS  JOINTS 


357 


the  neutralization  of  this  acid,  or  it  is  due  to  the  combina- 
tion of  all  these  factors  acting  through  their  influence  on 
the  ductless  glands. 

It  is  important  also  that  while  this  disease  is  very 
common  among  the  Cebidas  it  has  never  been  found  among 
the  macaques.  This  may  be  due  to  the  fact  that,  owing  to 
the  storage  sacs  in  the  mouth  of  the  macaques,  more  food 
proportional  to  body  weight  is  consumed,  or  there  may 
be  an  essential  difference  in  the  basal  metabolism  of  the 
families  and  individuals.  All  the  factors  enumerated  do 
tax  the  metabolic  resources  of  the  body  and  depress  the 
functions  of  the  endocrine  glands.  Only  detailed  and 
accurate  quantitative  studies  of  normal  metabolism  and 
the  effect  of  alterations  of  it  on  the  ductless  glands  will 
give  a  more  definite  answer  to  the  problem. 

This  work  indicates  clearly  the  alterations  to  be  made 
in  the  diets  to  meet  the  requirements  of  the  Cebidae  and 
is  to  be  followed  by  investigations  along  similar  lines 
for  other  families. 


Table 
A  List  of  the  Orders  Exhibiting  Definite  j 

17. 
jesions 

of  Osteomalacia  and  Rachitis. 

Osteomalacia 

Rachitis 

Primntp'S                                                                                    .... 

29 
4 
3 
2 
5 
1 

3 
9 
2 
12 

44 
35 

10 

Tipmiirps?                                                                                 

2 

8 

1 

3 

10 

34 

Psittaci      

Galli                             

79 

34=113 

Having  discussed  the  nature  of  these  diseases  and 
some  of  the  factors  in  their  causation,  analysis  of  their 
distribution  may  be  appropriately  added.  The  accom- 
panying list  (Table  17)  illustrates  the  orders  in  which  the 


358   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

two  diseases  have  been  found.  Veterinarians  are  familiar 
with  systemic  osseous  diseases  in  all  the  domesticated 
herbivores,  but  Hutyra  and  Marek  note  them  as  un- 
common in  dogs  and  birds.  Among  the  Primates,  osteo- 
malacia occurs  almost  exclusively  in  New  World  monkeys, 
Cebidae  and  Hapalidae,  whereas  rickets  is  much  more 
common  among  macaques  (CercopithecidaB).  Eight  of 
the  ten  cases  of  rickets  in  monkeys  seem  to  have  arrived 
at  the  Garden  vnth  e\ddences  of  this  disease.  Half  of  the 
cases  were  arrested,  or  at  least  not  florid,  when  the  beast 
came  to  autopsy.  All  of  the  osteomalacic  lemurs  belonged 
to  the  ring-tailed  species,  born  in  the  Garden  and  dying 
at  ages  from  three  to  seven  years.  The  cases  of  rickets 
among  the  Camivora  were  four  Felidae,  three  Canidae 
and  one  Procyonidae  while  all  the  osteomalacia  cases 
were  in  the  last  family.  Six  of  the  eight  cases  among 
the  rodents  affected  squirrels.  The  large  number  of  cases 
of  rickets  among  the  marsupials  is  due  to  a  litter  of 
small  opossums  thrown  by  an  apparently  healthy  mother 
and  dying  in  from  six  weeks  to  three  months. 

The  avian  varieties  w^hich  show  the  most  definite 
osteomalacic  changes  are  the  pigeons  and  pheasants,  mth 
the  parrakeets  presenting  nearly  as  characteristic  lesions. 
Birds  when  affected  with  this  disease,  may  come  to 
autopsy  in  fairly  good  plumage  and  wdthout  any  very 
marked  emaciation.  This  is  remarkable,  for  when  the 
cresta  sterni  is  palpated  tliis  ridge  may  sometimes  be 
bent  enough  laterally  to  touch  the  aim  sterni.  How  the 
bird  can  sit  upon  a  perch  when  it  is  possible  to  bend  the 
femora  almost  double,  is  difficult  to  understand.  De- 
formity is  by  no  means  so  frank  as  in  mammals  although 
periosteal  overgrowth  may  be  quite  marked  at  times. 
Anemia  is  undoubted  in  nearly  every  instance,  the  pallor 
of  the  muscles  seeming  to  be  as  great  as  if  the  specimen 
were  intentionally  bled  to  death.  \ 


THE  SKELETON  AND  ITS  JOINTS  359 

Osteitis  Deformans. 

Dr.  Corson- White  was  fortunate  enough,  during  the 
course  of  her  work  upon  osteomalacia  of  monkeys,  to 
detect  a  specimen  which,  did  not  show  the  usual  excessive 
excretion  of  calcium  but  on  the  other  hand  retained  this 
element  and  evinced  alkali  hunger.  The  general  appear- 
ance of  the  specimen  was  similar  to  that  of  monkeys 
having  osteomalacia  but  at  autopsy  a  definite  picture  of 
Paget 's  disease  or  osteitis  deformans  was  discovered. 
This  led  to  a  search  for  cases  in  the  literature  and  to  the 
following  study,  which  I  paraphrase  and  condense  from 
Doctor  Corson-White's  notes. 

C  Osteitis  deformans  is  a  chronic  constitutional  affection 
characterized  by  the  absorption  of  compact  bone,  cliiefly 
in  the  cranium  and  long  bones,  and  the  laying  down  of 
fibro-osteoid  tissue  in  such  an  excess  as  to  enlarge  the 
affected  bones.  This  material,  which  is  soft  and  cuts 
with  reasonable  ease,  has  calcareous  matter  in  it  as  shown 
by  Rontgen-ray  examination.  Paget  described  it  in  a 
classical  article  in  1876(3)  since  which  time  the  reported 
cases  have  mounted  to  three  hundred  and  fifty.  Because 
it  has  only  been  recognized  in  its  best  developed  stages, 
it  may  be  that  early  mild  or  arrested  cases  have  been 
overlooked.  Judging  by  the  instances  claimed  to  have 
been  found  in  museum  collections  of  bones,  it  is  probably 
an  affection  dating  to  antiquity.  So  far  these  remarks 
apply  only  to  man  but  in  lower  animals  the  reports  are 
very  few  and  those  are  not  available  in  the  original.  The 
abstracts  and  references  show  considerable  confusion. 
The  names  osteitis  deformans,  osteoporosis,  osteitis 
fibrocystica.,  osteodystrophia  deformans  and  osteosar- 
coma, leontiasis  ossei,  etc.,  are  used  almost  interchange- 
ably. ■  In  1901  Barthelemy  (4)  described  a  condition 
(Maladie  du  Son)  in  horses  in  which  there  was  a  marked 
enlargement  of  the  head  and  of  the  epiphyses  of  the  long 

(3)  Med.  Chir.  Trans.,  Vol.  60,  37,  1877. 

(4)  These  de  Lyon,  1901. 


360  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

bones.  His  cases  were  more  allied  to  osteitis  fibrosa 
cystica.  Paget 's  disease  always  attacks  the  diaphyses 
of  the  bones  and  not  the  epiphyses.  Goldman  (5)  de- 
scribed typical  examples  of  this  condition  in  fowls.  Jost 
(6),  in  one  commmiication,  desciibed  a  case  in  a  horse 
which  he  says  was  identical  with  that  condition  described 
by  Paget  as  osteitis  deformans  and  by  Virchow  as  leontia- 
sis  ossei ;  he  also  refers  to  similar  cases  in  goats  and  mon- 
keys. Rossweg(7)  found  it  in  goats.  In  wild  animals  the 
only  suggestive  article  found  was  by  Jost  but  the  descrip- 
tion was  probably  of  an  osteoporosis  and  a  craniosclero- 
sis  which  occurred  in  a  young  lion  and  a  monkey.  All  the 
communications  deal  mth  either  domesticated  animals  or 
those  in  captivity. 

The  etiology  of  this  condition  is  as  obscure  to-day  as 
it  was  at  the  time  of  Paget 's  first  description.  Prince 
thought  it  might  be  due  to  a  defect  in  some  peripheral 
nerve  or  nerve  centre  or  to  a  tract  degeneration.  Cases 
have  been  reported  in  conjunction  with  a  myelitis.  There 
has  been  however  little  on  which  to  base  these  supposi- 
tions. Paget  felt  that  the  process  was  at  least  upon  an 
inilammatory  basis  and  deduced  this  from  the  enlarge- 
ment and  the  excessive  production  of  an  imperfectly 
developed  structure  with  increased  blood  supply.  Many 
felt  that  rickets,  osteomalacia  and  osteitis  deformans 
were  all  manifestations  of  the  same  disease.  A  bacterial 
cause  was  proposed  by  Arcangelli  who  claimed  the  dis- 
covery of  diplococci  and  improvement  from  a  vaccine. 
Lancereaux(8)  and  Eichards  felt  that  focal  infection 
played  a  profound  role  in  the  etiology.  However  all  other 
observers  fail  to  isolate  an  organism  from  the  bones  or 
to   get  improvement  from  removal   of  infectious  foci. 

(5)  Verein  Freibilrger  Aerzte,  May,  1902. 

(6)  Arch.  f.  Wiss.  u.  Prak.  TierhJc,  Vol.  36,  652,  1910,  and  Vol.  39, 
164,  1913. 

(7)  Vet.  Med.  Inaug.  Diss.  Giessen,  1913. 

(8)  Traite  d' Anatomie  Path.,  1883. 


THE  SKELETON  AND  ITS  JOINTS  361 

Heredity  has  been  held  responsible  in  seven  per  cent,  of 
the  cases  in  human  beings. 

That  some  inflammatory  factor  is  partly  responsible 
seems  plausible  when  one  considers  the  active  growth  of 
fibroceUular  tissue  in  the  endo-  and  periosteum.  The 
more  interesting  theories  go  back  to  perversions  of 
internal  secretions,  pituitary,  parathyroid  etc.,  (Macallum 
&  Vogtlein).  Higbee  and  Ellis  (9)  say  in  relation  to  the 
neurotrophic  theory  that  if  the  neurotropliic  mechanism 
governs  metabolism  and  is  influenced  by  the  activity  of  the 
ductless  glands,  there  is  considerable  likelihood  that  its 
disturbance  may  possibly  be  found  to  be  the  cause. 

Da  Costa(lO)  believed  the  disease  to  be  a  disorder  of 
bone  metabolism  probably  dependent  on  the  absence  or 
perversion  of  some  internal  secretion.  There  is  much 
evidence  on  hand  to  indicate  that  disorders  of  the  ductless 
glands  do  influence  bone  metabolism,  and  changes  in  these 
glands  have  been  reported  in  cases  of  Paget's  disease, 
although  the  findings  and  lesions  have  been  far  from  uni- 
form or  distinctive  or  even  confined  to  one  gland.  Eight 
cases  were  reported  as  possibly  due  to  a  hypothyroid  con- 
dition; pituitary  changes  were  found  in  three;  adrenal 
changes  in  one;  parathyroid  reported  missing  in  two; 
three  had  sclerotic  thyroids.  Many  case  reports  make  no 
mention  whatever  of  the  glands  of  internal  secretion. 

Da  Oosta  interprets  the  retention  of  calcium,  phos- 
phorus and  magnesium,  with  the  sulphur  loss  found  in 
these  cases,  as  indicating  a  stimulated  osseous  or  osseoid 
formation  accompanying  the  resorption  of  a  highly  sul- 
phurized organic  matrix.  In  the  course  of  this  calcifica- 
tion procedure  we  suppose  a  certain  quota  of  the  sulphur 
of  the  matrix  is  replaced  by  other  elements,  a  process 
which  must  entail  retention  of  calcium,  phosphorus  and 
magnesium  and  increased  elimination  of  sulphur.  He 
shows  the  close  parallelism  between  the  mineral  metabo- 

(9)   Jour.  Med.  Res.,  Vol.  24,  43,  1911. 
(10)   Publ.  Jefferson  Med.  College,  Vol.  6,  1,  1915. 

24 


362   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

lism  of  a  gro\ving  boy,  a  case  after  parathyroidectomy  and 
a  case  of  osteitis  deformans,  and  suggests  that  this  de- 
pends in  some  way  either  on  the  absence  or  perversion  of 
some  internal  secretion,  possibly  of  the  parathyroids, 
which  controls  calcium  exchange  in  the  body.  Substances 
from  some  cause  arise  which  have  the  power  to  abstract 
calcium  from  the  body  tissues,  the  abstraction  of  these 
salts  being  the  first  step  in  the  production  of  the  disease. 

The  example  which  is  reported  in  full  was  the  first  to 
be  encountered  in  our  5,365  autopsies  but  shortly  after 
this  series  was  concluded  two  more  came  to  autopsy  and 
Dr.  Corson- Wliite  's  studies  were  made  to  embrace  these. 

A  reddish  woolly  monkey  {Lagothrix  infumatus) 
received  November  25,  1919,  was  a  particularly  active 
specimen  and  as  far  as  we  could  determine  a  perfectly 
healthy  adult  animal.  He  passed  the  tuberculin  test  and 
was  placed  on  exhibition.  In  April,  1920,  he  was  first 
reported  as  crippled  and  was  removed  to  the  laboratory  in 
June,  1920.  At  that  time  the  long  bones  of  the  legs  and 
arms  were  bowed  anteriorly  and  laterally,  the  degree  of 
curvature  making  the  hands  and  feet  seem  disconnected. 
The  monkey,  could  stand  but  made  no  voluntary  effort  to 
do  so.  There  was  evidently  some  pain  although  it  could 
not  have  been  at  all  severe.  He  resented  handling, 
especially  of  his  arms  and  legs.  The  head  was  rounded, 
resembling  that  of  a  baby,  and  the  eyes  were  protuberant 
suggesting  an  exophthalmos.  The  maxillary  bones  were 
so  excessively  thick  that  the  mouth  could  not  close  and 
the  monkey  drooled  saliva.  His  blood  on  admission  to  the 
infirmary  in  May  was  — Hg  eighty-nine  per  cent.,  R.  B.  C. 
4,370,000,  W.  B.'  C.  5,800;  one  week  before  death  it  was 
Hg  fifty-four  per  cent.,  R.  B.  C.  2,860,000,  W.  B.  C.  6,000. 
Routine  urine  examination  showed  a  constant  trace  of 
albumin,  hyaline  and  granular  casts.  There  was  at  no 
time  a  Bence-Jones  protein  reaction,  excess  of  indican, 
indol,  or  diacetic  acid.  He  had  a  constant  slight  diarrhoea 
with  some  flatulence,  the  semi-fluid,  constantly  acid  feces 


Fig.  46. — OSTEITIS  DEFORM.^N'S.  SKELETON  SHOWING  GENERAL  THICKENING  OF  ALL  BONES, 
BUT  ESPECIALLY  OF  THE  SKILL.  JAWS  AND  LONG  BONES.  NOTE  THICKNESS  OF  CALVARIUM. 
10  mm.:  NORMAL  IS  ABOUT  3-4  mm.  THE  HUMERUS  IS  TWICE  NOR.MAL  SIZE.  THE  WIDTH  OF 
THE  ULNAR  CORTEX  IS  SHOWN  BY  A  TRANSVERSE  SECTION  NEAR  THE  ELBOW  JOINT.  REDDISH 
WOOLLY  MONKEY   (LAGOTHRI.X  INFUMATUS). 


THE  SKELETON  AND  ITS  JOINTS 


363 


presenting  a  preponderance  of  Gram-positive  coccoid 
organisms.  His  appetite  was  fair  and  he  showed  marked 
craving  for  lime,  eating  plaster  from  the  walls  when  he 
could  get  it.  Because  of  this  desire  for  lime  he  was  given 
a  salt  mixture  to  see  if  it  would  have  any  effect  on  the 
bone  condition.     The  mixture  used  was  the  following : 


NaCl  0.874  grams 

KCl  0.548  grams 

CaH(PO)  H2O  3.608  grams 


Ca  lactate  0.386  grams 
Mg  citrate  0.848  grams 
K  citrate     1.953  grams 


This  mixture  the  animal  ate  with  avidity  and  seemed  more 
comfortable ;  other  than  that  no  change  was  noted. 

An  inorganic  metabolism  examination  was  attempted 
on  the  ordinary  diet  and  the  diet  plus  the  salt  mixture. 
Under  the  circumstances  at  our  disposal  this  was  not 
entirely  accurate  but  showed  such  marked  variation  from 
the  control  animals  examined — normal  and  osteo- 
malacic— and  such  marked  correspondence  on  the  four 
separate  four-day  periods  of  each  intake  that  it  seemed 
acceptable.  The  result  of  this  investigation  on  the  first 
series  of  four  four-day  periods  was : 


Intake 

Output 

Total 

Feces 

Urine 

0.0280 
0.0640 
0.1540 
0.1440 

0.014 
0.034 
0.027 
0.002 

0.0022 
0.0123 
0.0430 
0.1680 

0.0162  =  0.0118  retention 

Magnesium 

Phosphorus 

Sulphur 

0.0463  =  0.0177  retention 
0.07     =0.084    retention 
0.17     =0.026    loss 

The  result  on  the  higher  salt  content  was  practically  the 
same,  the  retention  being  in  proportion  slightly  less.  The 
diet  for  these  small  monkeys  is  two  apples,  two  bananas, 
six  small  sweet  potatoes,  with  a  lump  of  boiled  rice  about 
the  size  of  an  egg.  The  content  of  this  diet  has  been 
found  very  low  in  calcium,  phosphorus,  sodium,  chlorine 
and  iron,  while  potassium,  magnesium  and  sulphur  were 
high.  To  this  diet  lime  water  was  added  to  increase  its 
inorganic  content. 


364   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

This  Reddish  Woolly  Monkey  died  August  20,  1920, 
and  was  immediately  posted.  There  was  marked  thicken- 
ing of  the  frontal,  occipital  and  parietal  bones,  upper  and 
lower  jaws.  The  increase  in  the  size  of  the  alveolar 
margins  prevented  the  closing  of  the  mouth;  only  the  last 
four  teeth  could  be  brought  into  apposition.  The  enlarge- 
ment of  the  mandibles  reduced  the  capacity  of  the  mouth 
cavity.  The  skull  while  decidedly  thickened  did  not  en- 
large at  the  expense  of  the  cranial  cavity.  There  was  a 
cervical  and  dorsal  kyphosis.  The  chest  was  increased 
anteroposteriorly  and  contracted  laterally.  The  long 
bones  were  thick,  bulky  and  deformed.  (Fig.  46.) 

A  Black  Spider  Monkey  {Ateles  ater)  showed  a  gen- 
eral hyperplasia  of  the  whole  shaft  of  the  long  bones.  She 
was  much  deformed  by  curvatures  and  swellings  of  the 
skeleton — head  enlarged,  face  deformed  by  the  swelling  of 
the  upper  and  lower  alveolar  processes,  jaws  do  not  close 
and  the  palatal  bones  were  flattened,  skull  irregularly 
thickened,  elastic  but  not  soft,  slight  subperiosteal  growth. 
Thickening  of  the  long  bones  was  largely  due  to  subperios- 
teal growth;  section  of  the  ulna  showed  a  subperiosteal 
osseoid  layer  surrounding  the  old  shaft.  This  tissue 
seemed  to  be  very  poor  in  lime  salts,  cutting  without  any 
grit.  Marrow  cavity  was  filled  with  a  fairly  firm,  deep 
red  marrow  which  did  not  bleed  on  section.  The  third 
monkey,  a  Brown  cebus  {Cehus  fatuellus)  showed  exactly 
the  same  general  picture  but  was  less  severely  affected 
than  the  other  two. 

*'  The  more  minute  study  of  the  bones  of  these 
monkeys  shows  a  variety  of  pictures  while  preserving  one 
general  form.  The  skull  was  smooth,  mottled  by  irreg- 
ularly placed  areas  of  congestion ;  it  was  asymmetrically 
thickened;  differentiation  between  cortex  and  diploe, 
internal  and  external  tables  was  lost;  calcareous  matter 
was  absorbed  and  the  resultant  bone  was  soft  elastic  and 
porous ;  lacunae  enlarged  and  lined  with  bone  corpuscles 
and  giant  cells.     Other  areas  show  more  dense  bone,  the 


THE  SKELETON  AND  ITS  JOINTS  365 

reparative  processes  being  more  active  in  that  the  lamel- 
lae are  wide  and  the  vascular  spaces  narrower.  As  a 
rule  the  compact  bone  is  absorbed,  the  Haversian  canals 
are  more  or  less  confluent  and  there  is  generally  a  marked 
increase  of  newly  formed  osseoid  tissue.  The  ossifying 
periosteitis  obliterates  the  depressions  for  the  cranial 
arteries  and  the  sutures.  The  skull  cap  becomes  finely 
porous,  cancellous  and  even  cavernous.  The  spaces  are 
filled  with  a  soft,  red  marrow-like  material.  The  bones  at 
the  base  of  the  skull  are  much  less  involved;  the  peri- 
cranium, dura  and  brain  are  normal. 

**  The  long  bones  show  interlacing  narrow  strands 
w^hich  are  in  some*  regions  wide,  in  others  narrow  or  thin 
as  in  spongy  bone.  Large  irregular  cavities  are  present 
and  there  is  a  disappearance  of  the  compact  bone  and  an 
encroachment  on  the  medullary  canal  of  a  relatively  dense 
new  bone  with  small  irregular  trabeculae  surrounded  by 
osteoblasts  and  a  fibrous  connective  tissue  which  fills  the 
outer  trabecular  spaces.  The  new  bone  is  often  both 
subperiosteal  and  subendosteal,  the  latter  often  gaining 
on  the  former.  It  is  always  soft  and  irregularly  calcified. 
The  general  arrangement  of  the  strands  in  the  deep  layers 
is  longitudinal  but  in  the  subperiosteal  bone  they  are  very 
irregular  and  almost  at  right  angles  with  the  central 
strands.  In  this  new  osseoid  tissue  cysts  are  frequent, 
varying  in  size  from  very  small  to  rather  large  cavities 
filled  with  a  cloudy  gelatinous  material.  The  picture  here 
is  very  like  osteitis  fibrosa  cystica.  Endosteal  cells  pro- 
liferate and  may  fill  up  the  marrow  spaces  so  that  solid 
masses  of  fibrous  tissue  result.  Frequently  the  osteoid 
material  shows  fibrillae.  The  compact  bone  may  be  irreg- 
ular with  well  marked  Haversian  systems.  Toward  the 
periosteum  the  cells  may  become  scanty,  the  bone  dense  in 
structure  while  toward  the  interior  the  cells  are  more 
numerous  and  in  the  more  cancellous  portions,  the 
trabeculae  become  slender  and  far  apart ;  here  they  may 


366   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

be  covered  by  a  single  row  of  osteoblastic  cells.  The 
intertrabecular  spaces  are  large,  irregular  and  filled  with 
a  delicate  alveolar  tissue  containing  only  a  minimal 
number  of  normal  bone  marrow  cells,  large  capillaries  and 
no  giant  cells.  Periosteum  may  be  of  usual  thickness; 
the  bone  immediately  beneath  is  spongy.  The  sclerosis 
of  the  bone  in  its  densest  areas  is  entirely  due  to  the  ossi- 
fication of  spindle  cells  which  have  remained  in  the  place 
of  the  original  marrow  of  the  bone.  As  a  whole  the  bone 
is  nowhere  normal  in  amount  or  proportion  but  the  small 
Haversian  systems  are  properly  made,  the  abnormality 
being  chiefly  due  to  cellular  and  fibrous  growth  around  the 
large  lamellae  which  at  times  is  normal  in  amount  but 
usually  much  in  excess.  In  places  this  consists  wholly  of 
fibroblasts,  at  others  of  giant  and  round  cells  very  sug- 
gestive of  sarcoma.  All  histologists  apparently  agree 
that  Paget 's  disease  starts  as  a  resorption  of  already 
calcified  bone." 

These  data  seem  to  supply  ample  evidence  that  the 
autopsy  diagnosis  of  Paget 's  osteitis  deformans  was  cor- 
rect. While  the  anatomy  and  course  and  chemical  changes 
presented  by  these  monkeys  do  not  settle  the  causation  of 
the  disease,  they  offer  very  definite  suggestions  which 
Dr.  Corson-White  summarizes  in  the  following  cau- 
tious conclusions. 

'*  Many  of  the  cases  of  Paget 's  disease  first  came 
under  the  observation  for  fractures,  accidents  common  in 
osteomalacia  but  very  rare  in  developed  cases  of  osteitis 
deformans.  Early  cases  all  presented  diarrhoea,  which 
was  present  in  all  the  early  human  cases  seen,  and  in  all 
the  cases  reported  in  monkeys.  This  symptom  was  men- 
tioned in  fourteen  of  the  cases  from  the  literature.  It  was 
also  a  constant  symptom  in  primate  osteomalacia.  The 
diet  of  these  monkeys  was  exceedingly  low  in  those  sub- 
stances essential  to  bone  development,  and  Sherman  has 
shown  that  the  calcium  balance  is  regulated  to  a  certain 


THE  SKELETON  AND  ITS  JOINTS  367 

extent  by  the  calcium  ingested,  and  that  when  the  diet  was 
poor  in  this  element,  the  output  materially  exceeds  the 
intake,  a  fact  which  is  immediately  changed  where  the 
animal  is  put  on  a  diet  high  in  calcium.  So  far  as  we  could 
find  there  are  no  studies  on  the  mineral  metabolism  of 
beginning  cases  of  Paget 's  disease.  It  seems  possible 
from  the  osteomalacic  animals  previously  studied,  that  the 
low  mineral  and  otherwise  faulty  content  of  the  diet  might 
so  disturb  the  chemical  equilibrium  directly,  through  the 
neurotrophic  mechanism  or  through  the  perversion  of  the 
ductless  glands,  that  the  mere  addition  of  lime  water 
might  entirely  change  the  pathological  picture.  This  is  in 
accord  with  the  histology.  The  initial  histological  pic- 
ture is  always  resorption  of  bone,  a  general  decalcification 
which  later  presents  an  irregular  proliferation.  The 
disease  then  progresses  along  different  lines  ending  as 
osteitis  fibrosa  cystica,  Paget 's  or  Von  Recklinghausen's 
diseases,  etc.,  dependent  upon  the  strength  of  the  repara- 
tive stimulus  and  the  organism  upon  which  it  acts. 

''  These  cases  are  of  interest  (1)  because  they  are  typi- 
cal examples  of  Paget 's  disease  as  it  has  been  described 
in  man  both  clinically  and  pathologically,  (2)  because  the 
disease  shows  the  same  general  type  of  inorganic  metab- 
olism that  was  exliibited  in  man,  (3)  because  of  the  alkali 
hunger  shown  by  one  monkey,  and  by  two  human  cases,  a 
hunger  which  was  severe,  which  preceded  the  deformity 
and  disappeared  after  the  deformity  was  established, 
(4)  because  the  disease  developed  in  animals  fed  on  a  diet 
insufficient  in  its  inorganic  and  vitamine  content  to  which 
an  excess  of  calcium  was  added. 

**  From  this  study  it  seems  possible  that  Paget 's 
disease  may  be  just  one  stage  in  a  deficiency  disease,  a 
reparative  response  through  a  neurotrophic  mechanism 
or  through  the  perversion  of  the  glands  governing  calcium 
metabolism  wliich  has  been  perverted  by  an  improperly 
balanced  diet." 


368   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Tumors. 

Neoplastic  diseases  of  the  bones  have  yet  to  be  classi- 
fied to  everyone 's  satisfaction.  All  gradations  of  hyper- 
plasia of  osteogenetic  cells  and  fibres  and  of  the  marrow 
elements,  from  simple  inflammation  to  true  sarcoma,  are 
recognized.  As  one  reviews  a  large  series  of  lesions, 
clearly  defined  types  may  be  fomid,  but  there  are  transi- 
tion stages  to  which  an  exact  name  is  difficult  or 
impossible  to  apply.  Nor  does  the  pathological  diagnosis 
always  fit  Avith  the  clinical  course.  Thus,  for  example,  the 
giant-cell  tumor  of  bones  looks  malignant,  and  is  not,  and 
its  structure  may  be  simulated  in  such  diseases  as  fibrous 
osteitis  and  Paget 's  disease.  As  we  have  seen  in  the  dis- 
cussion of  the  latter  of  these  two,  abnormalities  of  fibre 
and  cell  gro^\i;h  simulate  neoplasms  very  closely.  In 
addition  it  might  be  mentioned  here  that  actinomycosis 
may  produce  bony  growths  resembling  sarcoma.  When 
osteitis  deformans  affects  the  facial  bones  especially,  it 
has  been  called  leontiasis  ossium,  and  it  is  then  a  more 
nodular,  tumor-forming  process,  the  enlargement  con- 
sisting of  fleshy  masses  occupying  the  whole  bone,  but 
especially  the  marrow  cavity.  Histologically  the  lesion  is 
fibrocellular,  frequently  with  numerous  giant  cells; 
accumulations  of  small  sarcoma-like  alveoli  may  be  found. 
We  have  encountered  four  cases  among  common  opos- 
sums {Didelphys  virginiana)  and  one  in  an  Isabelline 
Gazelle  {Gazella  Isabella)  and,  because  of  the  localization 
and  fleshy  consistency  of  the  tumor,  we  have  called  them 
osteosarcomata  for  descriptive  purpose,  but  not  for 
classification  among  neoplasms  (where  they  will  not  be 
found).  Two  of  the  opossums  had  osteoporosis  and 
gelatinous  marrow  in  the  ribs.  All  these  animals  had  been 
in  the  exhibition  under  a  year,  and  as  far  as  known  are 
not  related.  Figure  47  shows  the  gross  character,  while 
the  following  is  the  description  from  one  protocol. 


Fig.  47. — EXAMPLES  OK  LOCAL  O.STEOMAIA  RESE.VIBLING  OS  lEOSARCOMA  AND  FIBROUS 
OSTEITIS;   THEY   PROBABLY   BELONG  TO  THE  DISEASE  KNOWN   AS   LEONTIASIS   OSSIUiVl. 

A  AND  B,  OPOSSUMS  (DIDELPHYS  VIRGINIANA).  C,  ISABELLINE  GAZELLE  (GAZELLA 
ISABELLA). 


Fig.  48. — FIBRO-OSTEOMA,  A  LOCAL  SINGLE  TUMOR.  OK  THE  UPPER  JAW.  THIS  DID  NOT 
RESEMBLE  THE  CHANGES  IN  ACTINOMYCOSIS.  BUT  THE  INFECTION  WAS  NEVERTHELESS  EX- 
CLUDED   BY   BACTERIOLOGICAL  SEARCH.      ISABELLINE  GAZELLE  (GAZELLA  ISABELLA). 


THE  SKELETON  AND  ITS  JOINTS  369 

Common  Opossum  {Didelphys  virginiana)  9.  Osteosarcoma  of  al- 
veolus, rarefying  osteitis  deformans  of  skull,  hypertrophy  of  thyroid, 
acute  catarrhal  enteritis,  acute  hyperplasia  of  spleen.  About  the  middle 
of  both  lower  rami  and  involving  the  posterior  half  of  each  upper 
maxilla  is  a  uniform  elliptical  growth  apparently  emanating  from 
alveolus.  Teeth  not  loose,  but  can  be  moved  in  tumor  to  be  described. 
On  section  a  white  glistening  homogeneous  growth  is  seen  apparently 
originating  in  the  body  of  the  alveolus  and  around  the  teeth.  The  shaft 
of  bone  is  soft  and  easily  broken.  What  remains  of  marrow  is  irregu- 
larly injected.  In  upper  jaw  there  is  a  distinct  porosis  of  facial  bones; 
they  and  the  enclosed  sinuses  are  deeply  injected.  Lower  four  ribs  on 
both  sides  show  distinct  nodulations  of  pale  color  along  a  bluish 
bone.  All  ribs  are  very  soft  and  section  shows  osteoporosis  of  shaft 
Avith  injected  marrow  and  distinct  cartilaginous  periosteal  bone  forma- 
tion. The  skull  is  everywhere  soft  and  the  bone  is  apparently  in- 
creased in  thickness,  rich  in  blood,  but  porotic.  Rest  of  skeleton  seems 
well  calcified.  Microscopic  section  of  tumor  shows  practically  the 
same  picture.  Bone  is  nowhere  normal  in  amount  and  proportion  but 
the  Haversian  systems  seem  properly  made,  the  abnormality  consisting 
chiefly  of  cellular  and  fibrous  growth  around  larger  lamellae,  which  at 
times  is  normal  in  amount  but  usually  much  in  excess.  In  places  this 
consists  wholly  of  fibroblasts,  at  others  of  giant  and  round  cells  very 
suggestive  of  sarcoma;  indeed  all  areas  must  be  called  giant-cell  sar- 
coma. There  is  an  attempt  to  lay  down  osteoid  tissue  at  places  par- 
ticularly beneath  periosteum.  The  giant  cells  are  in  great  numbers 
and  some  seem  osteoclastic.  As  the  lamellee  disappear  young  connective 
tissue  seems  to  take  their  place  but  giant  cells  do  not  remain  numerous 
at  such  places.  Despite  its  atypical  nature  it  must  probably  be  looked 
upon  as  an  osteosarcoma.  The  tooth  socket  is  not  much  involved  save 
for  hyalinization  of  root  matrix  immediately  about  dentinal  zone. 

In  another  Isabelline  Gazelle  (Gazella  isabella)  there 
was  a  fibro-osteoma  localized  to  one  side  of  the  superior 
maxilla  (Fig.  48) ;  this  has  been  included  among  the 
tumors,  while  the  above  mentioned  cases  have  not  been 
so  grouped. 

True  osteosarcoma  seems  not  to  have  occurred.  One 
tumor  was  seen  upon  the  wing  of  a  Cuvier's  Toucan 
{Rhamphastos  cuvieri)  which  was  formerly  diagnosed  as 
sarcoma,  but  later  examination  reveals  some  giant  cells 
in  arrangement  suggestive  of  tuberculous  osteitis ;  since 
this  is  the  only  case  and  not  unequivocally  a  tumor  its 
record  is  hardly  warranted.  Osteomata  of  the  hard 
variety  have  been  seen  on  the  ribs  of  a  pigeon  and  a 


370   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

pheasant  as  small  rounded  compact  well  outlined  tumors. 
It  is  thought  that  they  represent  products  of  healing  after 
osteomalacia  or  rickets.  An  osteochondroma  growing 
from  the  nasal  cartilage  was  found  in  a  caracal  {Felis 
caracal),  a  fibro-osteoma  was  found  on  the  vertebra 
and  clavicle  of  a  Beechj^'s  spermophile  {Citellus  gram- 
mums  heecheyi)  and  a  fibroma  occurred  on  the  clavicle 
of  a  lesser  snow  goose;  {Chen  hyperboreus  hyperboreus). 
The  only  other  tumor  from  a  bone  was  an  endothelioma 
from  the  periosteum  of  the  clavicle  in  a  moorhen  (GaWi- 
nula  chloropus) .  It  corresponds  to  the  usual  idea  of  this 
tumor.  It  probably  caused  death  by  cachexia,  and  by  its 
size,  interference  Avith  respiration.  'The  only  secondary 
tumor  was  a  metastasis  in  the  tibia  from  a  spindle  cell 
sarcoma  of  the  kidney  in  a  scaly  ground  dove  {Scarda- 
pella  squamosa). 

The  Muscles. 

The  skeletal  muscles  of  the  wild  animals  of  our  collec- 
tion have  been  quite  free  of  pathological  lesions  such  as 
atrophies  and  dystrophies  and  indeed  seem  relatively 
seldom  affected  by  disease.  Occasionally  hyalinization 
will  accompany  infectious  disease  or  local  suppurations 
will  spread  into  the  muscles.  Much  more  often  filaria, 
sarcocystis,  flukes  and  larval  insects  will  be  found  resi- 
dent within  or  between  muscle  bundles;  this  vdW  be 
discussed  at  a  later  time.  Six  tumors  have  been  found, 
three  of  which  certainly  developed  in  a  muscle,  while  for 
a  fourth  case  no  primary  growth  was  discovered.  This 
last  one,  to  dispose  of  it  at  once  because  of  its  peculiarity, 
was  an  adenocarcinoma  found  as  a  firm,  conglomerate, 
encapsulated  mass  in  the  sheath  of  the  gluteal  muscles  of 
a  waltzing  mouse  {Mus  ivagneri  rotans).  No  other  growth 
was  discovered  although  it  must  be  admitted  the  body  was 
not  exhaustively  searched  for  some  tiny  nodule  to  which 
primary  focus  this  muscle  mass  could  have  been  sec- 
ondary.    That  metastases  may  be  larger  than  original 


THE  SKELETON  AND  ITS  JOINTS  371 

growths  is  well  knowm.  The  gross  diagnosis  was  sarcoma. 
If  this  be  an  original  tumor  it  might  be  explained  as 
arising  from  ectopic  mammary  tissue. 

The  five  other  tumors  were  sarcomata,  one  of  large 
cells  almost  syncytial  in  size,  shape  and  number  of  nuclei, 
two  definite  spindle  cell  growths  and  two  of  fibrosarcoma 
type.  The  first  occurred  in  an  all-green  parrakeet 
{Brotogerys  tirica),  the  second  in  an  undulated  grass 
parrakeet  (Melopsittacus  undulatus),  the  third  in  a  larger 
Egyptian  gevhiWe  ){Gerhillus  pyramidum),  the  fourth  in 
a  white-footed  mouse  (Peromyscus  leucopus) ,  (smd  the 
last  in  a  bean  goose  {Anser  fabalis). 


SECTION    XIII 

THE  CENTRAL  NERVOUS  SYSTEM  AND 
THE  SPECIAL  SENSES 

Diseases  of  this  system  and  its  specialized  end  organs 
of  sense  in  the  eye,  ear  and  nose  are  recognized  by 
veterinarians  as  occurring  among  domesticated  animals, 
in  which  however  they  are  by  no  means  so  common  as  in 
^/ human  beings.  Because  of  the  natural  reserve  of  wild 
beasts  and  because  their  habits  and  manners  are  not  so 
familiar  to  the  observer,  it  would  seem  that  clinical  evi- 
dences of  disease  of  the  nervous  system  are  rarest  among 
them.  The  brain,  cord  and  nerves  of  our  specimens  have 
not  been  studied  with  the  minuteness  accorded  to  the 
examination  of  other  viscera  since  we  have  not  had  the 
persoimel  to  devote  the  time  to  this  really  colossal  under- 
taking, our  work  with  these  tissues  being  directed  toward 
the  accumulation  and  preservation  of  apparently  normal 
brains  from  every  species;  there  are  now  some  nine 
hundred  brains  on  the  shelves.  However,  whenever  the 
history  of  the  animal  before  death  has  suggested  that 
disturbance  in  the  nerve  organs  might  exist,  they  have 
been  dissected  grossly  and  examined  microscopically. 
Had  we  subjected  all  our  material  to  microscopic  study 
it  is  quite  possible  that  we  might  have  discovered  more 
lesions.  This  is  still  possible  by  reason  of  our  gross 
material  and  autopsy  protocols. 

We  have  been  fortunate  in  having  Dr.  W.  B. 
Cadwalader,  Secretary  of  the  Society,  and  Dr.  J.  H.  W. 
Rhein  ^vith  their  broad  experience  in  neurology,  take 
interest  in  this  phase  of  the  subject  and  study  our 
material.  The  number  of  cases  in  which  definite  lesions 
have  been  found  is  surprisingly  small,  so  that  no  general- 
izations can  be  attempted  except  perhaps  in  a  negative 
sense.    After  the  study  of  thirty-nine  brains  and  cords 

372 


THE  CENTRAL  NERVOUS  SYSTEM  373 

from  animals  in  whose  history  some  suggestion  of 
nervous  system  disease  existed,  Doctor  Cadwalader  could 
find  abnormalities  in  only  twenty-two  cases,  mostly  how- 
ever of  a  very  indefinite  character.  This  observer 
further  points  out  that  in  his  experience  with  our 
material,  his  observations  of  our  animals  and  a  review  of 
the  literature,  he  is  unable  to  find  satisfactory  instances 
of  the  so-called  system  or  tract  diseases  such  as  tabes 
and  lateral  sclerosis.  This  he  suggests  is  due  to  the 
absence  of  arterial  sclerosis  in  the  lower  types  of  brain, 
intimating  further  that  perhaps  the  relation  of  syphilis 
toi  human  arteriosclerosis  and  degenerative  nervous  sys- 
tem disease  is  thereby  strengthened.  These  facts  being 
true  it  is  not  astonishing  that  massive  cerebral  apoplexy 
and  cerebral  softening  do  not  occur (1).  The  lesions  that 
have  been  found  were  either  definitely  infectious,  as 
encephalitis  or  acute  septicemia,  tuberculosis,  poliomye- 
litis, meningitis,  or  the  minute  hemorrhages  and  vague 
granularities  or  vacuolizations  of  nerve  cells  in  toxemic 
conditions.  The  material  being  limited  and  broad  state- 
ments being  impossible,  the  important  cases  will  be  cited 
individually  or  in  small  groups. 

Convulsions. 

Before  entering  upon  the  special  subjects,  a  word 
might  be  added  concerning  the  clinical  evidences  of 
neurologic  conditions  as  seen  in  the  Garden.  Perhaps  the 
most  common  and  certainly  the  most  definite  clinical  sign 
of  nervous  disturbance  is  the  convulsion.  General  spasms 
or  fits  are  fairly  common  among  the  carnivores  and 
monkeys,  in  the  former  most  often  associated  with 
intestinal  parasitism,  in  the  latter  with  no  especial 
relationship  unless  it  be  renal  disease.  The  convulsions 
have  been  general,  with  and  without  the  preservation  of 

( 1 )  We  have  observed  later,  however,  one  case  with  very  definite 
degenerative  arterial  disease;  it  will  be  discussd  under  the  affections  of 
the  eye  since  the  most  definite  and  indeed  only  lasting  sign  of  trouble  was 
amblyopia,  the  attack  being  ushered  in  by  a  nondescript  fit. 


374   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

consciousness.  No  cases  of  focal  or  Jacksonian  spasms 
are  recorded  although  we  shall  cite  two  instances  of 
tuberculoma  reasonably  near  the  motor  area  in  the 
cerebrum.  The  exact  cause  of  many  cases  cannot  be  set 
down  since  intracranial  disease  does  not  often  exist. 

Intoxication  from  chemical  products  of  disturbed 
digestion  or  from  worms  themselves  is  the  usual  expla- 
nation of  spasms  associated  with  intestinal  parasitism; 
if  this  be  true,  intoxication  in  the  absence  of  demon- 
strable parasites  may  also  be  the  cause  in  certain  cases 
of  enteritis.  However  we  have  seen  fits  when  enteritis, 
parasites  and  renal  disease  were  not  found.  If  these 
were  true  epilepsy,  they  are  instances  of  perhaps  the 
rarest  disease  of  animals,  which  I  do  not  presume 
to  diagnose. 

Convulsive  seizures  in  herbivorous  animals  are  ex- 
ceedingly rare  although  I  have  seen  clonic  movements 
of  a  spasmodic  character  in  antelopes  and  deer  shortly 
before  death  from  gastroenteritis.  Ataxia  and  incoordi- 
nation are  much  more  common.  ,  Birds,  notably  parrots 
and  soft-billed  insectivorous  varieties,  are  not  uncom- 
monly afflicted  with  fits  but  as  they  are  rarely  observed 
except  by  the  keeper  the  exact  nature  is  difficult  to 
describe.  Those  seen  by  the  writer  have  been  of 
two  kinds. 

The  first  and  more  common  consists  of  falling  from 
the  perch  in  a  dazed  and  stiff  condition,  with  dilated  fixed 
eyes,  stiffened  and  spread-out  legs  and  wings.  Eecovery 
follows  shortly  and  the  bird  resumes  its  perch  either  in 
excitement,  or  slowly  and  uncertainly,  perhaps  to  have 
another  attack  in  a  few  minutes.  These  cases,  in  the 
few  instances  in  which  they  could  be  followed,  were  due 
to  faulty  feeding  and  enteritis  and  showed  either  nothing 
or  a  mild  congestion  of  the  brain.  The  other  variety  of 
fit  is  epileptiform,  a  rapidly  developing  clonic  spasm  of 
all  parts  of  the  body  with  a  tendency  to  opisthotonos.  In 
one  case  of  this  character,  a  parrot,  no  lesions  were  found 


THE  CENTRAL  NERVOUS  SYSTEM  375 

ill  the  brain,  an  enteritis  existing  however.  Another 
case  concerned  a  pet  Indian  Shama  I  had  at  my  home. 
He  had  been  doing  well  and  singing  loudly,  until  one 
evening  he  was  allowed  to  remain  in  a  tobacco-smoke- 
filled  room  whereupon  next  morning  he  stopped  eating 
and  singing.  Later  that  day  the  clonic  form  of  con- 
\'Tilsions  appeared,  growing  worse  for  thirty-six  hours  or 
until  death.  At  autopsy  no  food  was  found  in  the 
alimentary  tract.  The  brain  and  cord  were  congested 
grossly,  while  minutely,  perivascular  hemorrhages  and 
marked  vacuolization  and  diffuseness  of  staining  were 
found  in  ganglion  cells  of  the  bulb,  pons,  anterior  spinal 
horns  and  in  the  pyramidal  cells  of  the  cerebellum.  Hap- 
pening so  promptly  after  exposure  to  tobacco  smoke, 
when  the  bird  was  doing  well,  I  venture  to  associate 
the  two. } 

Ataxia. 

Incoordination  and  ataxia  are  so  often  observed  and 
under  so  many  conditions  that  it  is  well  nigh  impossible  in 
any  individual  case  to  give  an  adequate  explanation 
before  death.  They  are  in  all  probability  the  expression 
of  sickness  and  nothing  more  in  the  vast  majority  of 
instances.  When  they  are  observed  in  such  cases  as  the 
tyromata  of  the  cerebrum  or  in  certain  of  the  migulates, 
they  may  mean  something  definite.  In  this  latter  order 
and  to  a  less  extent  in  carnivores,  one  frequently  sees 
weakness  and  uncertainty  of  gait  in  the  hind-quarters,  the 
legs  being  usually  coordinate  but  tending  to  give  way 
under  the  weight  of  the  body. 

From  a  study  of  veterinary  literature  and  our  own 
material  it  would  seem  that  this  may  have  many  explana- 
tions. In  the  first  place,  it  may  simply  indicate  weakness 
expressing  itself  in  the  heaviest  part  of  the  body,  the 
animal  inclining^  its  femora  forward  to  assist  in  support- 
ing the  hea^^  abdomen.  It  may  be  an  expression  of 
abdominal  pain,  the  recti  becoming  rigid  and  the  quadri- 


376   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

ceps  of  the  thigh  participating  in  the  protection  of  the 
belly.  Almost  any  of  the  intra-abdominal  conditions, 
gastroenteritis,  mesenteric  thrombosis,  peritonitis,  or 
diseases  of  the  psoas  muscle  and  lumbar  vertebra,  might 
occasion  this  attempt  at  support.  Disease  of  gluteal 
muscles,  as  hemoglobinuric  fever,  may  produce  a  palsy  of 
the  whole  pelvic  girdle  with  weakness  of  the  hind  legs. 
There  may  be  associated  with  the  weakness  of  the  hind 
legs  a  humped-up  condition  of  the  lumbar  spine  and 
retraction  of  the  abdomen,  sometimes  called  ' '  tucked  in ; " 
in  two  definite  cases  of  this  last  sort  we  have  found  renal 
pelvic  stones  and  once  intestinal  sand.  Some  instances 
are  undoubtedly  due  to  meningomyelitis  or  to  poliomye- 
litis and  at  the  place  for  this  subject  a  few  cases  will  be 
discussed.  Meningitis  has  not  been  found  in  the  ungu- 
lates showing  this  weakness.  There  have  been  however 
cases  of  ataxia  in  the  hind  legs  of  deer  and  antelopes, 
which  did  not  have  a  ready  explanation  fitting  in  with  the 
foregoing.  Two  of  these  we  thought  might  be  due  to 
certain  grasses  in  the  enclosures  and  have  changed  the 
exhibition  spaces.  No  conclusion  can  be  drawn  from  this 
as  yet.  No  enterocolic  disease  could  be  found  nor  any 
lesion  of  the  sciatic  nerve  and  lumbar  enlargement  of  the 
cord.  We  have  however  discovered  sciatic  neuritis  in  a 
case  like  hemoglobinuric  fever  in  a  Burchell  's  zebra.  The 
history  of  the  animal  is  similar  to  that  of  this  disease  in 
domestic  animals  in  so  far  as  symptomatology  is  con- 
cerned ;  in  so  far  as  confinement  in  a  stall  is  concerned  no 
data  is  at  hand  but  death  occurred  on  December  26th  in 
the  zebra  house  whereas  he  had  been  accustomed  to  go 
out  into  the  yard  all  summer  and  autumn. 

Meningitis. 

The  coverings  of  the  brain  and  cord  have  not  been  the 
seat  of  the  well  known  acute  inflammations  seen  in 
domestic  horses  and  cattle.  Eleven  instances  of  disease 
in  the  meninges  are  recorded  but,  with  very  few  excep- 


THE  CENTRAL  NERVOUS  SYSTEM  377 

tions,  have  been  accompanied  by  other  lesions  offering  a 
ready  etiological  explanation.  These  cases  are  however 
not  very  instructive  except  perhaps  three  in  monkeys 
where  the  meningitis  seemed  to  be  secondary  to  gastro- 
enterocolitis.  In  one  case  a  colon  bacillus  was  apparently 
responsible,  in  a  second  no  bacteriology  was  undertaken 
and  a  third  was  too  rotten  for  the  results  to  be  dependable. 
A  focus  of  infection  aside  from  the  intestinal  area  could 
not  be  found.  The  only  noteworthy  finding  was  the 
scantiness  of  the  cerebrospinal  fluid  and  the  almost 
exclusive  subpial  exudate;  these  facts  would  seem  to 
strengthen  the  thought  that  the  virus  came  through  the 
blood  stream.  Another  case  was  due  to  extension,  through 
the  temporal  bone  to  the  lateral  sinus,  of  a  necrotizing 
process  beginning  in  the  buccal  muscles  or  parotid  gland; 
the  necrosis  bacillus  and  a  host  of  Gram-positive  cocci 
were  found.  A  Canadian  porcupine  suffered  with  a 
mucopurulent  nasopharyngitis  which  involved  the  deep 
sinuses,  the  middle  ear  and  the  temporal  bone;  smears 
from  the  pus  over  the  corresponding  cerebral  hemisphere 
and  from  the  nasal  pus  showed  pneumococcus  forms ;  the 
lungs  were  not  affected.  The  llama  which  showed  the 
intracapsular  fracture  of  the  femur  (page  344)  had  also 
hemorrhage  into  a  fibrinous  exudate  in  the  mastoid  cells 
with  deep  opaque  congestion  and  edema  ot  the  pia  above 
the  petrous  portion  of  the  temporal  bone.  Decomposition 
precluded  satisfactory  bacteriology  but  it  is  suggested 
that  probably  injury  in  falling  started  a  hemorrhage  in 
the  ear  upon  which  a  secondary  infection  was  implanted. 
Wliat  seems  a  true  meningitis  secondary  to  otitis  media 
and  mastoid  suppuration  was  seen  in  a  marmoset. 

A  case  of  the  well  kno'^vn  but  obscure  condition  known 
as  chronic  productive  pachymeningitis  was  observed  in 
a  badger.  Although  it  cannot  be  explained  it  is  cited  as 
a  matter  of  record  and  interest. 

American  Badger  (Taxidea  taxus).    Pachymeningitis  externa.    The 
dura  is  fast  to  the  skull  and  cannot  be  removed.    Scattered  irregularly 
25 


378   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

over  the  entire  inner  surface  of  the  skull  are  pale  pinhead  sized  hard 
nodules.  It  is  impossible  to  tell  if  they  are  in  the  dura  or  the  bone. 
The  brain  shows  engorged  vessels  but  is  otherwise  negative. 

Two  instances  of  hemorrhagic  pachymeningitis  as- 
sociated with  cretinism  were  seen  in  wolf  cubs.  The  f  ol- 
lomng  notes  illustrate  both  cases. 

American  Gray  Wolf  {Canis  mexicanus) .  Cretinism.  Hemorrhagic 
external  pachymeningitis  with  craniotabes.  Upon  removing  the  calva- 
rium  a  marked  craniotabes  of  the  under  surface  is  found  and  with 
it  a  deep  red  and  purple  staining  of  outside  of  dura  and  inner  table 
of  skull.  These  changes  are  most  marked  along  the  longitudinal  sinus 
at  internal  occipital  protuberance  and  along  left  parietal  region.  Dura 
on  left  side  is  distinctly  congested.  This  is  also  true  of  pia.  The 
brain  and  its  base  seem  normal. 

Cyst  of  the  brain.  A  sooty  mangabey  {Cercocebus 
fidiginosus)  had  been  in  the  Garden  for  about  four 
months  and  was  apparently  an  adult  normal  animal.  It 
died  rather  suddenly  after  a  distinct  con\ailsion  with  semi- 
consciousness. At  autopsy  in  addition  to  a  nephritis,  a 
large  cyst  was  found  to  occupy  the  posterior  third  of  the 
left  hemisphere.  Its  walls  were  composed  of  a  thin  (one- 
sixteenth  inch)  rim  of  cerebral  substance  and  the 
meninges;  its  contents  were  clear.  The  notes  do  not 
record  any  examination  for  parasites.  This  monkey 
showed  no  localizing  signs. 

Tuberculosis. 

Gross  tuberculous  lesions  have  been  found  in  the  brain 
in  several  specimens  with  generalized  disease  but  only 
two  cases  are  of  special  interest.  A  Rhesus  macaque 
(Macaciis  rhesus)  suffering  with  generalized  but  chiefly 
lymphatic  tuberculosis,  showed  a  large  plaque  on  the 
external  surface  of  the  dura  over  the  vertex  where  it  was 
adherent  to  a  yellow,  fairly  firm  nodule  about  ten  mm. 
across.  This  nodule  was  deeply  imbedded  in  the  brain 
substance,  barely  projecting  above  the  surface,  generally 
spherical  and  not  encapsulated.  There  was  no  peripheral 
reactive  zone.  The  meninges  were  not  altered  anywhere 
except  as  above.     The  blood  vessels  were  not  especially 


THE  CENTRAL  NERVOUS  SYSTEM  379 

congested.  The  pia  arachnoid  contained  no  excess  of 
fluid  but  the  summits  of  the  convolutions  were  flattened. 
The  mass  was  located  in  the  posterior  frontal  convolution, 
near  the  longitudinal  fissure,  occupying  nearly  its  whole 
breadth  and  penetrating  about  one  centimetre.  It  did 
not  enter  the  fornicate  gyrus.  There  was  a  completely 
degenerated  core  about  two  mm.  across.  The  adjacent 
bone  was  beginning  to  erode.  No  localizing  signs 
were  reported. 

The  other  case,  that  of  a  young  Drill  baboon  (Papio 
leucopJiceus) ,  was  studied  with  Dr.  J.  H.  W.  Rhein  and 
can  be  reported  in  the  following  condensed  notes. 

The  baboon  appeared  to  be  perfectly  well  until  October  18,  1906, 
when  some  lameness  in  the  anterior  and  posterior  extremities  on  the 
right  side  was  observed.  This  gradually  increased,  and  was  associated 
with  general  convulsions.  On  November  30th,  I  made  an  examination 
and  found  the  following  condition :  The  right  upper  and  lower  extremi- 
ties were  weaker  than  on  the  left  side.  On  the  left  side  the  power  seemed 
to  be  fair.  He  was  able  to  hold  on  to  an  iron  bar  with  the  fingers  of 
the  upper  and  lower  extremities  on  the  right  side  but  in  A\dthdrawing 
the  bar  it  was  not  difficult  to  overcome  his  grasp  and  the  power  on  this 
side  was  distinctly  less  than  on  the  left.  The  movements  of  the  right 
arm  were  someAvhat  ataxic,  as  observ^ed  when  he  made  efforts  to  grasp 
the  bar.  The  knee  jerks  were  increased  on  both  sides  and  appeared 
to  be  equally  so.  There  was  no  evidence  of  facial  palsy.  He  moved 
both  sides  of  the  face  equally  well  at  times  when  he  expressed  anger 
or  fear  in  the  facial  expression.  The  tongue  seemed  to  be  retracted 
equally  well  on  both  sides.  Tests  for  hemianopsia  Avere,  of  course,  un- 
satisfactory, but  he  seemed  to  recognize  readily  the  approach  of  the 
iron  bar  from  both  sides.  There  was  no  disturbance  of  the  rectal  or 
bladder  functions,  although  at  autopsy  the  bladder  was  full.  Death 
occurred  on  December  2,  1906.  At  the  autopsy  the  brain  and  spinal 
cord,  with  the  other  organs  were  examined.  The  dura  was  adherent 
to  the  left  side  of  the  brain,  in  the  prefrontal  region,  in  the  upper 
third  and  when  the  brain  was  removed  it  was  observed  that  an  area 
of  softening  lay  beneath  this  point.  A  small  caseous  mass  was  also 
observed  at  the  base  of  the  right  lung,  and  beneath  the  diaphragm  on 
the  right  side  was  a  large  abscess,  partly  involving  the  liver.  The 
tubercle  bacillus  was  found  in  the  pus  removed  from  the  area  of  soften- 
ing beneath  the  left  cortex.  A  study  of  the  brain  revealed  the  presence 
of  three  foci  of  softening.  The  largest  one  was  situated  in  the  left 
hemisphere  in  the  prefrontal  region,  and  extended  from  just  beneath 
the  cortex  in  the  upper  third  of  the  region,  downward  almost  to  the 
base  of  the  brain.     This  area  was  cylindrical  in  shape  and  measured 


380   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

2.5  cm.  in  its  greatest  diameter.  The  area  of  softening  consisted  of 
caseated  material  and  pus,  in  which  the  tubercle  bacillus  was  found 
present.  The  second  area  of  softening  was  found  on  the  right  side 
of  the  brain,  much  smaller  in  extent  and  measuring  1.5  cm.  diameter. 
There  was  no  pus  present  in  this  area,  but  it  consisted  of  a  circumscribed 
mass  of  caseous  material.  The  apex  of  this  area  of  softening  Avas  just 
beneath  the  cortex  in  the  prefrontal  region  on  the  right  side,  and  in 
the  removal  of  the  brain  the  cortex  was  torn  just  above  this  area  of 
caseation.  This  point  was  .5  cm.  in  front  of  the  central  fissure  and 
about  1  cm.  beloAV  the  superior  surface  of  the  brain.  A  third  area  of 
softening  was  observed  posteriorly,  in  the  white  substance,  in  the 
parieto-occipital  region,  and  measured  about  6  mm.  in  diameter.  A 
study  of  the  sections  of  the  brain  shows  beautifully  the  extent  of  the 
destruction  of  the  brain  tissue.  The  optic  thalamus  and  the  lenticular 
nucleus,  and  the  posterior  limb  of  the  internal  capsule,  on  the  left 
side,  are  destroyed  in  part.  The  anterior  limb  of  the  internal  capsule 
in  one  section,  is  preserved,  notwithstanding  the  fact  that  the  optic 
thalamus  on  the  same  side  has  been  in  large  part  destroyed.  In  spite 
of  the  fact  that  the  posterior  limb  of  the  internal  capsule  has  been 
destroyed  at  some  levels,  it  is  interesting  to  note  that  the  degeneration 
of  the  pjTamidal  tracts  of  the  pons  and  medulla  on  the  opposite  side 
and  the  lateral  columns  of  the  spinal  cord  are  not  intensely — although 
distinctly — degenerated.  It  is  very  interesting  to  note  that  in  view 
of  the  severe  damage  to  the  posterior  limb  of  the  internal  capsule  on 
the  left  side,  there  was  not  more  paralysis,  for  it  will  be  remembered 
that  there  was  considerable  power  of  prehension  in  the  right  upper 
and  lower  extremities. 

Encephalomyelitis,  Poliomyelitis. 

While,  as  has  been  stated,  no  well  defined  cases  of  the 
recognized  meningocephalic  infectious  diseases  have  been 
observed  in  the  Garden,  there  have  been  several  animals 
in  whose  cord  and  brain  changes  were  found  comparable 
to  the  infectious  disease  known  in  man  as  poliomyelitis. 
On  several  occasions  since  this  disease  was  recognized  as 
occurring  in  epidemic  form  it  has  been  observed  that 
domestic  mammals  and  fowls  suffered  from  a  similar 
condition.  The  general  vicinity  of  Philadelphia  had  a  low 
grade  epidemic  among  children  during  the  years  from 
1907  to  1912  and  it  is  during  this  time  that  most  of  the 
cases  of  a  comparable  character  were  observed  among 
our  animals.  It  is  to  be  emphasized  that  attacks  were 
entirely  sporadic  and  the  cases  did  not  appear  to  bear  a 


THE  CENTRAL  NERVOUS  SYSTEM  381 

relation  to  one  another.  This  character  is  quite  in  accord 
with  certain  of  the  outbreaks  in  man.  However  we  can- 
not state  that  the  disease  is  exactly  the  same  as  seen  in 
the  human  being  for,  as  will  be  noted  in  the  cited  examples, 
all  the  pathological  features  were  not  fulfilled. 

It  is  not  always  possible  to  differentiate  between 
myelitic  disease  and  polyneuritis  of  man  or  animals. 
Studied  symptomatically  the  cases  in  our  records  which 
proved  to  have  degenerative  and  infiltrative  lesions  com- 
parable to  poliomyelitis  showed  gradual  but  progressive 
paralysis  expressed  by  inability  to  move  rather  than  dis- 
inclination— in  other  words  loss  of  power  rather  than 
restriction  because  of  pain.  None  of  the  animals  in 
which  poliomyelitis  was  demonstrated  have  exhibited  the 
ataxia  of  the  hind  legs  discussed  on  a  previous  page  nor 
have  we  found  myelitic  lesions  in  the  few  cords  from 
animals  suffering  with  this  weakness.  The  nearest  ap- 
proach to  a  cerebrospinal  explanation  for  weakness  and 
palsy  was  in  a  zebra  which  died  with  constipation,  acute 
nephritis,  and  hepatic  perilobular  fibrosis.  In  this  animal 
a  pronounced  subpial  mononuclear  infiltrate  was  ob- 
served, in  places  invohdng  the  superficial  parts  of  the 
cerebral  gray  matter,  especially  about  the  congested 
vessels  of  this  area.  This  condition  was  present  to  a 
slight  degree  in  the  cord.  There  was  then  a  low  grade 
meningoencephalitis  but  no  nerve  cell  changes.  So  far  as 
is  known  to  me  no  animal  showing  a  definite  local  paresis 
or  paraplegia  recovered  from  the  attack;  had  this 
occurred  we  might  have  observed  residual  palsies. 

In  so  far  as  lesions  are  concerned  they  are  perhaps 
best  illustrated  by  the  appended  cases,  but  since  even  in 
them  there  is  a  lack  of  uniformity,  it  may  be  well  to 
discuss  the  basic  changes  of  all.  The  outstanding 
abnormality  in  the  microscopic  anatomy  is  the  richness 
of  small  mononuclear  cells  beneath  the  pia,  both  spinal 
and  cerebral,  around  the  smaller  blood  vessels  and  to  a 
lesser  extent  around  the  multipolar  cells  of  the  gray 


382   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

matter.  These  do  not  present  the  dense  colonization  often 
seen  in  the  acute  cases  of  infantile  paralysis  in  man  but 
are  prominent  in  comparison  to  normal  nervous  tissue. 
Hemorrhages  or  at  least  small  groups  of  erythrocytes 
outside  of  blood  vessels  are  seen  here  and  there.  Vacuoli- 
zation of  ganglion  cells  is  variable,  being  prominent  in 
some,  trifling  or  absent  in  others.  G-Ual  proliferation  is 
often  quite  marked,  and  in  one  case  to  be  cited  seems  the 
prominent  lesion. 

The  animals  in  which  meningopoliomyelitis  has  been 
found  were  three  monkeys,  two  Canadian  lynx,  a  bear  and 
a  raccoon;  about  a  score  of  cords  from  other  animals 
with  some  kind  of  palsy  have  been  studied  microscopically 
mthout  discovering  it.  The  following  cases  illustrate  our 
material.  The  only  instance  of  two  cases  in  close  relation 
concerns  the  lynx  {Fells  canadensis).  They  occupied  the 
same  cage  and  died  twelve  days  apart.  No  symptoms 
were  recorded  until  a  few  days  before  death  when  a 
general  paralysis  appeared,  deepening  to  completeness  on 
the  day  of  death.  No  case  occurred  in  neighboring  cages. 
Doctor  Rhein  studied  all  these  cases,  and  his  notes  are 
used  for  these  records.  Portions  of  the  lumbar  and  cervi- 
cal enlargements  and  of  the  thoracic  regions  of  the  cord 
were  stained  with  hemalum  and  acid  fuchsin  and  Avith 
thionin.  The  pia  was  slightly  infiltrated.  There  was 
some  cellular  infiltration  of  the  anterior  septum,  and  the 
vessels  here  showed  an  increase  in  the  nuclei  of  the  walls 
and  a  slight  perivascular  infiltration.  The  pial  infiltra- 
tion seemed  to  be  equally  distributed  in  the  entire  circum- 
ference of  the  cord,  although  perhaps  a  little  more 
marked  over  the  anterior  and  posterior  septa.  The  ves- 
sels of  the  gray  matter  were  congested  and  the  walls  of 
the  vessels  in  most  part  showed  a  proliferation  of  the 
nuclei.  There  were  a  few  small  hemorrhages  into  the 
gray  matter,  probably  agonal.  As  compared  with  the 
human  cord  and  the  cords  of  monkeys,  antelopes  and 
dogs,  there  was  an  unusually  large  number  of  gUa  nuclei, 


THE  CENTRAL  NERVOUS  SYSTEM    383 

which,  if  found  in  the  human  cord,  would  be  looked  upon 
as  a  proliferation  process.  In  some  cases  these  nuclei 
were  heaped  together  in  masses,  and  were  evidently 
pathological.  There  was  also,  about  the  ganglion  cells, 
some  pericellular  round  cell  infiltration,  and  this  was 
more  marked  around  a  few  cells  which  were  almost 
entirely  destroyed.  The  ganglion  cells  themselves  were 
swollen.  Some  showed  eccentric  nuclei,  and  many  of  them 
stained  poorly,  while  one  or  two  showed  distinct  vacuo- 
lization. In  one  field  a  ganglion  cell  was  partly  destroyed 
by  a  recent  hemorrhage.  There  were,  however,  a  number 
of  cells  which  appeared  normal.  This  process  seemed  to 
be  fairly  distinct  in  the  lumbar  and  cervical  enlargements, 
but  was  not  clearly  demonstrated  in  the  sections  from 
the  dorsal  region.  The  cellular  infiltration  of  the  horns 
was  evidently  not  leucocytic,  but  presented  the  appear- 
ance of  a  connective  tissue  proliferation.  Although  these 
are  not  the  exact  lesions  found  in  poliomyelitis  in  the 
human  animal,  they  are  at  least  suggestive  of  the  same 
process  since  the  infiltrating  cells  are  of  the  lymphatic  or 
connective  tissue  types.  There  is  no  acute  inflammatory 
leucocytic  infiltrate. 

A  weeper  cebus  (Cebiis  capucinus)  ever  since  he  was 
received  acted  in  such  a  peculiar  manner,  seeming  to  have 
only  partial  control  of  his  movements,  that  he  was  known 
as  the  "Crazy  Monkey."  There  was  no  history  of  illness 
before  death.  Pathological  diagnosis:  Chronic  entero- 
colitis, chronic  adhesive  pericarditis,  early  interstitial 
change  in  kidney,  edema  of  lungs,  meningitis  and  polio- 
myelitis. The  pia  of  the  paracentral  cortex  was 
thickened  and  was  the  seat  of  a  round  cell  infiltration  of 
moderate  degree,  the  cells  being  of  the  mononuclear  type. 
The  blood  vessels  of  the  cortex  were  congested  and  the 
nuclei  of  the  walls  were  increased.  The  round  cell  infil- 
tration of  the  pia  had  in  some  places  extended  into 
the  cortical  layers.  The  pia  surrounding  the  medulla 
oblongata  was  also  the  seat  of  a  slight  round  cell  infil- 


384   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

tration.  The  pia  of  the  spinal  cord,  however,  did  not 
show  any  cellular  infiltration.  The  cells  of  the  anterior 
horns  of  the  spinal  gray  matter  were  extensively  diseased, 
being  swollen  in  places,  some  surrounded  by  a  glia  pro- 
liferation and  many  mth  marked  vacuolization. 

A  common  raccoon  {Procyon  lotor)  was  observed  in 
the  laboratory  to  have  complete  paralysis  of  the  anterior 
and  partial  paralysis  of  the  posterior  extremities.  This 
latter  was  almost  complete  in  muscles  controlling  the 
feet,  while  the  thigh  and  hip  muscles  showed  some  irregu- 
lar incoordinate  movements.  Respiration  shallow  but 
regular.  History  shows  that  the  power  of  the  extremities 
began  to  fail  about  a  month  before  death  and  was  abso- 
lutely lost  in  the  fore  extremities  three  days  before  the 
animal  was  killed.  Diagnosis :  Poliomyelitis.  Examina- 
tion of  the  central  nervous  system  showed  the  presence  of 
marked  round  cell  infiltration  of  pia  of  cortex  and  of 
spinal  cord,  more  particularly  in  the  lower  thoracic  and 
lumbar  regions.  Ganglion  cells  in  the  lumbar  region  were 
markedly  diseased.  There  were  numerous  old  and  fresh 
hemorrhages  and  a  moderate  degree  of  round  cell  infiltra- 
tion in  the  anterior  horns.  Two  young  of  this  animal, 
bom  three  months  before  its  death,  showed  weakness  and 
gradual  increasing  paralysis  of  their  extremities  begin- 
ning when  three  months  old  (that  is  at  the  time  of  the 
death  of  their  mother),  and  lasting  until  their  death,  one 
in  the  seventh  and  one  in  the  eighth  month  of  life.  These 
were  found  not  to  have  changes  in  the  central  nervous 
system,  but  there  was  sufficient  rachitis  to  account  for 
this  paralysis. 

The  only  tumor  of  the  central  nervous  system  found 
among  these  animals  occurred  in  an  Undulated  Grass 
Parrakeet  {Melopsittacus  undulatus).  The  gross  notes 
are  very  vague  but  the  microscopy  is  suggestive  of  a 
glioma.  The  growi;h  in  the  brain  consists  of  large  irregu- 
lar masses  of  large  cells  mth  vesicular  nuclei  and  pale 
homogeneous    protoplasm.      *' Scattered   between   these 


THE  CENTRAL  NERVOUS  SYSTEM  385 

accumulations  are  irregular  strands  of  spindle  cells,  with 
spindle-shaped  nuclei,  taking  the  hematoxylin  very 
deeply.  The  supporting  tissue  is  almost  without  cells, 
taking  the  eosin  faintly,  and  is  quite  loosely  arranged. 
No  fibrils  are  seen  among  the  cells.  The  blood  vessels  are 
congested,  and  at  one  place  there  is  a  small  hemorrhage. 
The  vessel  walls  are  the  same  as  the  rest  of  the  connective 
tissue.  There  is  a  slightly  atypical  metastasis  in 
the  liver." 

A  case,  the  identity  of  which  is  still  undecided,  was 
observed  in  a  Green  Monkey  {Cercopithecus  callitri- 
chus) ;  it  may  belong  among  the  gliomata  or  glioses. 
There  was  in  the  middle  of  this  monkey's  cerebrum  a  gray 
area  about  3x2x1  cm.  with  a  softened  centre,  the  more 
solid  parts  being  found  under  the  microscope  to  consist 
of  glia  tissue,  blood  vessels  and  degenerated  cells.  No 
true  gliomatous  formations  could  be  discovered.  Because 
of  the  iiidefiiiiteness  of  the  lesion,  it  is  not  included  in 
the  tumors  or  inflammations.  Clinically  the  effect  of  the 
change  was  to  cause  blindness  and  ataxia  but  motor 
power  was  not  greatly  impaired. 

A  very  small  number  of  tumors  of  the  brain  in  wild 
animals  is  on  record  in  the  English  and  German  litera- 
ture, perhaps  the  most  interesting  being  what  resembles 
in  description  a  subdural  neurocytoma  reported  by 
Wilson  in  the  Proceedings  of  the  London  Zoological 
Society,  1908.  The  mass  was  separate  from  the  cere- 
bellum, but  had  hollowed  out  a  place  for  itself  in  this  part 
of  the  hind-brain. 

Beain  "Weights. 

The  policy  of  preserving  the  brain  of  all  species 
enables  us  to  record  in  the  accompanying  list  the  weights 
of  a  large  number  of  specimens.  In  order  that  the  figures 
may  have  a  representative  and  comparative  value  only 
those  are  given  where  the  total  body  weight  of  the  animal 
is  also  knowai.    The  specimens  were  removed  by  the  lab- 


386   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

oratory  staff,  most  of  them  by  one  person,  and  by  the 
same  technique.  All  brains  were  weighed  immediately 
upon  removal  from  the  body,  no  preservative  being  near 
the  organ.  The  brains  were  themselves  externally  nor- 
mal. Our  technician  is  skillful  in  removing  the  organ, 
practically  always  getting  the  pituitary-  body,  and  cutting 
off  the  brain  stem  at  the  foramen  magnum,  the  pia 
remaining  but  the  dura  removed.  Because  the  speci- 
mens were  taken,  in  practically  all  cases,  from  animals 
that  died  in  the  Park,  and  because  of  the  shortness  of  the 
list  (196),  it  seems  wise  not  to  attempt  conclusions  refer- 
able to  comparative  weights  of  the  different  orders  and 
families.  However,  the  data  seem  worthy  of  record 
because  it  is  doubtful  if  anywhere  one  can  find  so  many 
weights  taken  under  comparable  conditions  by  the  same 
personnel.  One  can  find  a  considerable  list  of  brain 
weights  and  values  in  many  publications  throughout  the 
literature,  notably  in  an  article  by  Ziehen  in  Bardeleben's 
Eandbuch  der  Anatomie  (Vol.  IV,  Abt.  Ill  363),  but  from 
no  single  source  are  there  so  many  varieties  or  so  long  a 
list.  Ziehen's  tables  are  compiled  from  the  literature 
and  therefore  represent  data  collected  under  different 
conditions,  many  of  which  were  probably  pathological. 
The  appended  figures  are  to  be  considered  as  raw 
material  collected  under  uniform  conditions. 

Examination  of  the  figures  bears  out  in  a  measure 
some  of  the  remarks  made  by  Ziehen,  notably  those 
which  indicate  that  between  large  and  small  varieties 
of  the  same  general  group,  the  smaller  has  the  greater 
brain  weight  value  and  that  the  youthful  animal  has  more 
brain  than  the  adult. 

The  brains  at  the  museum  are  fixed  in  saline-formalde- 
hyde— sufficient  strength  of  the  former  to  suspend  the 
organ  in  the  container  and  four  percentage  of  the  latter. 
When  fixation  is  complete,  as  indicated  by  density,  preser- 
vation is  done  in  one  per  cent,  formaldehyde,  the  organ, 
usually  bound  in  gauze,  being  laid  in  cotton.    A  list  of 


THE  CENTRAL  NERVOUS  SYSTEM  387 

important  references  is  added  to  the  weight  tables — some 
antedating  Ziehen's  articles,  but  principally  those  that 
have  appeared  since  the  publication  of  his  monograph. 

Principal  References  to  the  Relative  Weight  of  the  Brain 
Ziehen:    In  Bardelehen's  Haiidbuch  der  Anatomie  des  Menschen,  Vol.  4, 

pt.  3. 
Ziehen:  Handbuch  der  Anatomie  des  Nervensy stems,  Jena,  1903. 
Girabd:  Bulletin  de  V  Institut  Gen.  Psychologic,  Vol.  7,  p.  53. 
VON  BuscHAU:   Real  Encyclopedic  der  Gesamten  Heilkunde  3.  Aufl. 
VON  BuscHAU:  Neurologisches  Zentralbl.,  1897,  March. 
Brandt:  Bull,  de  la  8oc.  Imperial  des  Naturalistcs  de  Moscow,  1867,  40, 

pt.  2,  525. 
Mies:  Verhandl.  der  GeseUsch.  deutscher  Naturforsch  und  Arzte,  1898,  353. 
Dhere  et  Lapicque:  Arcliives  de  la  Physiologic,  October,  1898. 
Lapicque  et  Girard:   C.  R.  des  Sceances  de  I' Academic  de  Science,  Paris, 

1905,  140,  1057. 
Lapicque:  Bulletin  de  Museum  d'Histoire  Naturelle,  1909,  No.  7,  408. 
Lapicque:  Revue  du  Mois,  Paris,  April,  1908,  445. 
Lapicque:   Bulletin  et  Memoires  de  la  Societe  d' Anthropologic  de  Paris, 

1907,  5,  Vol.  8,  No.  3,  261. 
Lapicque:  Biologica,  Vol.  2,  1912,  p.  257. 
Funk:  Inaug.  Dissert.  Wurzburg,  1911. 
POYNTER:   Cerebral  Anthropology,  Lincoln,  1913. 
MoLLisoN:  Arch,  fiir  Anthropologic,  1914,  XIII,  388. 
HuLTGREN:  Das  Himgeuncht  des  Menschen,  Upsala,  1912. 
Weber:  Festschrift  fiir  Karl  Gegenbauer,  1898. 
Dubois:  Bulletin  de  la  Soc.  Anthropologic,  Paris,  1897,  337. 
DiTBOiS:   Archiv.  fiir  Anthropologic,  Vol.  25,  1898. 
Dubois:  Proc.  8ci.  K.  Acad.  Wet.,  Amsterdam,  1914,  16,  647. 
Dubois:   Zeitschrift  fiir  Morph.  und  Anthropologic,  1914,  Vol.  18,  323. 
Marchand:  Himgewicht  des  Menschen,  Leipsig,  1902. 
Rudolph:  Beitrdgc  zur  Path.  Anatomic,  Jena,  58,  1914,  48. 
Kraemer:   Mitt,  der  Deutsch.  Landwehrgesell,  29,  1914,  55. 


388   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 


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SECTION  XIII— PART  II 

THE  EYE 
Inflammations. 

Conjunctivitis  is  rather  rare  but  does  occur  in  all 
varieties  of  animals,  seldom  however,  to  the  extent  that 
the  specimen  has  to  be  sacrificed.  So  far  as  the  daily 
reports  can  be  relied  upon  we  have  had  no  epidemic 
inflammations  of  the  eye.  The  parrots  and  monkeys  are 
the  only  animals  that  can  be  handled  enough  for  treat- 
ment, and  in  them  the  applications  have  not  seemed  very 
efficacious.  Two  cases  of  conjunctivitis  and  iritis  have 
had  a  tuberculous  basis  and  two  other  specimens  have 
had  tuberculosis  in  the  eye.  Two  parrots  had,  in  asso- 
ciation with  generalized  tuberculosis,  semisolid  masses  in 
the  orbit  which  dislocated  the  bulb,  thickened  the  lids  and 
presented  as  yellowish  granulating  tumors  under  the 
conjunctiva;  a  Swainson's  long-tailed  jay  {Calocitta  for- 
mosa)  had  this  lesion  on  both  sides.  The  only  case  in 
a  mammal  concerned  an  Anubis  baboon  {Papio  anubis)  in 
which  one  eye  had  been  enucleated,  two  weeks  before 
death,  for  purulent  ophthalmitis.  The  other  eye  became 
affected  shortly  after  the  extraction  and  the  animal  was 
killed;  it  was  found  to  have  caseous  pneumonia.  The 
tuberculous  process  had  probably  begun  in  the  region  of 
the  optic  nerve  and  involved  the  whole  bulb.  Secondary 
infection  with  pyogenic  cocci  had  been  superimposed 
upon  the  original  process.  Keratomalacia,  encountered 
on  a  few  occasions,  will  be  discussed  by  Doctor  White. 

Iridocyclitis  was  found  in  a  white  browed  guan 
{Penelope  superciliaris)  the  notes  of  which  are  condensed 
as  follows : 

The  internal  organs  show  nothing  of  value  pathologically.  The 
right  eye  shows  moderate  conjunctivitis  and  a  marked  thickening 
of  the  cornea  with  complete  opacity.     The  lens  is  destroyed  and  the 

402 


THE  EYE  403 

retina  infiltrated  by  gelatinous  material.  Humors  are  watery,  non-suppu- 
rative,  but  the  fundal  portion  of  the  retina  shows  several  poorly  circum- 
scribed, yellowish  white,  gelatinous  collections.  Microscopical  section 
of  cornea  shows  replacement  of  normal  bundles  by  wavy  ones  inter- 
mixed with  small  numbers  of  nuclei.  These  are  never  of  inflammatory 
type,  but  always  of  connective  tissue  type.  Conjuctival  mucosa  defec- 
tive in  centre,  puckered  but  shows  no  subjacent  inflammatory  features. 
Ciliary  body  richly  infiltrated  by  Ijonphocytes  and  vessels  distinctly 
congested.  This  condition  extends  over  whole  anterior  surface  of  iris 
and  for  a  short  distance  over  posterior.  Sclera  shows  much  bone  for- 
mation. Chronic  interstitial  keratitis,  subacute  interstitial  iritis  and 
cyclitis. 

Cataract. 

Opacities  of  the  cornea  are  quite  common  among  our 
specimens,  most  often  due  we  believe,  to  local  trauma; 
ungulates  exhibit  them  more  than  other  varieties.  Cata- 
ractous  opacities  of  the  lens  are  frequently  observed  in 
senile  animals  but,  while  I  have  no  figures  for  the  state- 
ment, I  believe  they  are  not  as  common  among  our 
specimens  as  can  be  observed  in  domestic  horses  and 
dogs.  No  record  has  been  made  of  streaky  clouds  or  spots 
in  the  lens  but  only  of  complete  opacities.  There  are  three 
only,  an  aoudad  {Ovis  tragelaphus)  a  macaw  {Ara  macao) 
and  a  summer  duck  {Aix  sponsa)  the  last  of  which  alone 
is  interesting.  This  bird,  a  fully  developed  adult  female, 
was  killed  because  of  total  blindness  and  found  to  have  a 
low  grade  chronic  pancreatitis  and  a  bilateral  Morgag- 
nian cataract,  the  lens  capsule  containing  a  thin  cloudy 
fluid  with  the  nucleus  quite  freely  movable  in  it. 

Amblyopia. 

A  very  interesting  case  of  amblyopia  in  a  young 
monkey  was  studied  and  reported  by  Dr.  H.  M.  Langdon 
and  Doctor  Cadwalader  in  the  Journal  of  Comparative 
Pathology  and  Therapeutics,  Vol.  XXVIII,  Part  4. 
Because  of  its  unusual  character  and  careful  investiga- 
tion, the  report  is  reproduced  here  : 

Pigtailed  macaque  {Macactis  nemestrinus)  was  bom  June  9,  1913, 
in  the  monkey  house,  a  Avell  developed  baby.     He  thrived  and  was  as 


404  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

good  as  any  for  his  age.  He  was  never  known  to  have  anything  wrong 
with  him  until  on  the  morning  of  June  3,  1914,  when  he  was  found  on 
the  bottom  of  the  cage  in  the  monkey  house.  He  had  clenched  hands 
and  feet,  jaws  tightly  closed,  lips  drawn  back,  eyes  staring  and  glassy, 
with  convulsive  shaking  of  the  extremities.  At  intervals  he  would 
become  limp,  with  fists  still  clenched,  and  Avith  only  occasional  jerks  in 
the  extremities.  This  would  last  about  a  minute,  and  then  convulsive 
movements  would  be  resumed.  The  entire  ''fit"  lasted  about  ten 
minutes.  He  was  immediately  removed  from  the  large  cage  in  the 
monkej'  house  to  a  small  one  in  the  back  room  of  the  laboratory.  When 
put  in  the  small  cage  he  staggered  as  if  dazed,  and  groped  about 
apparently  blind.  He  never  recovered  his  sight  entirely,  but  at  times 
seemed  to  see  better  than  at  others.  He  was  not  seen  in  a  "  fit "  in  the 
laboratory.  On  June  24th,  a  small  piece  of  banana  was  offered  by  a 
person  who  stood  directly  in  the  sunlight.  The  monkey  came  to  the 
front  of  the  cage,  reached  out  and  grasped  very  firmly  the  thumb  of 
the  hand  holding  the  banana  but  did  not  take  the  banana  although  he 
very  plainly  wanted  it.  The  banana  was  thrown  into  the  cage,  hitting 
the  monkey  on  the  back.  He  turned  very  quickly,  then  smelled  over 
the  floor  of  the  cage  until  he  found  the  banana.  On  June  30th,  he 
was  examined  by  Doctor  Langdon  and  the  following  condition  was  noted : 

"  Pupils  react  to  the  light  of  the  ophthalmoscope.  Optic  discs  are 
normal.  Arteries  possibly  a  little  small.  No  other  fundus  changes." 
On  July  1,  a  cloudy  day,  he  was  laid  facing  a  window.  A  coat  sleeve 
was  laid  over  his  eyes  for  a  minute  and  then  quickly  removed.  His 
pupils  were  seen  to  react  slowly  but  distinctly  to  the  light.  His  gaze 
would  not  follow  a  finger  moved  in  front  of  his  eyes.  "When  put  back 
in  the  cage  he  climbed  up  on  the  wire  at  the  back  and  then  tried  to 
climb  the  plain  sheet-iron  side.  He  groped  and  felt  for  a  support  and 
then  fell.  This  he  did  several  times.  About  August  1,  when  the  eyes 
Avere  examined,  there  seemed  to  be  more  visual  perception  and  very  dis- 
tinctly prompter  pupillary  reflex,  which  condition  remained  about  the 
same  when  examined  October  1.  He  died  October  10,  1914,  of  a  com- 
pound fracture  of  the  right  femur  inflicted  by  a  monkey  in  the  ad- 
joining cage. 

At  autopsy  the  viscera  appeared  normal  throughout.  The  animal 
was  fairly  well  nourished.  There  was  about  5  cc.  clear,  pale  yelloAv  fluid 
under  the  dura.  It  escaped  upon  removal  of  the  brain.  There  were 
adhesions  of  the  dura  over  the  temporal  lobe  (inferior  surface),  poste- 
rior and  external  to  the  optic  tract,  so  firm  as  to  remove  some  periosteum 
and  superficial  bone.    Rest  of  dura  seemed  normal. 

Examination  of  the  brain.  Sections  were  made  from  different  parts 
of  the  cerebral  cortex,  all  of  which  were  more  or  less  alike.  There  was 
SAvelling  of  the  endothelium  of  the  pial  lymph  spaces,  with  some  separa- 
tion of  the  fibres  of  the  pia  itself  which  extended  into  the  sulci.  The 
perivascular  lymph  spaces  of  the  larger  arteries  of  the  cortex  Avere 
dilated,  and  the  adjacent  cerebral  tissue  was  edematous.  A  well  marked 
endothelial  swelling  and  hyperplasia  affected  a  number  of  the  arteries 


THE  EYE  405 

and  capillaries  producing  marked  general  or  nodular  thickening  in 
some  places.  Accompanying  these  hyperplastic  changes  there  was  a 
marked  calcification  of  some  of  the  arteries.  This  was  not  confined 
to  one  tunic,  but  in  some  instances  it  extended  almost  completely  through 
the  vessel  wall,  and  here  and  there  the  lumen  of  a  vessel  was  nearly 
obliterated.  The  main  features  were  endothelial  hyperplasia,  edema  of 
the  pia  and  of  the  subpial  cortex  with  some  calcification  of  the  vessels. 
It  was  perhaps  less  well  marked  in  the  occipital  lobes  than  in  other 
parts.  The  optic  nerve  and  other  portions  of  the  brain  appeared  to 
be  normal. 

Moon  Blindness. 

It  seems  also  profitable  to  repeat  here  a  report  Dr.  H. 
M.  Langdon  and  I  made  in  1911  upon  a  horse  with  periodic 
ophthalmia  or  '^moon  blindness,"  a  widespread  condi- 
tion and  one  upon  which  there  is  even  to-day  little  known 
and  much  contradictory  theorizing.  It  is  worthy  of 
record  that  Dr.  J.  H.  W.  Eyre  of  Guy's  Hospital,  had  a 
case  to  study  at  the  same  time  as  ours.  He  did  not  find 
the  protozoon-like  body  discussed  below,  but  laid  weight 
upon  the  isolation  of  St.  aureus,  an  organism  often  men- 
tioned in  the  literature  about  this  disease.  I  cite  the 
whole  report  since  our  publication  in  the  1911  Report  of 
this  Garden  seems  not  to  have  been  quoted  in  any  of  the 
reference  articles  on  ''Moon  blindness."  Those  who  are 
interested  in  the  clinical  and  pathological  sides  of  the 
question  will  find  a  good  summary  in  Veroff.  aus  der- 
Jahres.  Vet.  Berichten  der  beamt.  Tierdrzte  Preussens, 
1908,  and  the  bacteriology  of  the  equine  eye  by  Karsten, 
Inaug.  Disser.  Giessen,  1909. 

''During  the  latter  part  of  1909  and  first  part  of  1910 
we  had  a  horse  referred  to  us  suffering  with  recurrent 
ophthalmia  or  moon  blindness.  This  affection,  suggested 
by  its  name,  is  supposed  to  have  some  relation  to  the  lunar 
periods.  Some  points  in  our  work  showed  that  such  may 
be  the  case.  Attacks  appear  not  infrequently  at  the  time 
of  the  full  moon,  and  in  our  only  experimental  infection 
twenty-eight  days  elapsed  between  inoculation  and  a 
general  ocular  inflammation. 


406   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

''This  affection  manifests  itself  as  a  conjunctivitis  early 
in  the  attack,  but  rapidly  progresses  to  an  iridocyclitis  and 
lastly  to  a  panophthalmitis.  After  each  attack  the  ball 
is  smaller  until  it  is  so  shrunken  as  to  be  sightless  from 
chronic  thickening  and  opacities.  The  causation  is  not 
known.  The  disease  behaves  not  unlike  an  infectious  one, 
remaining  in  a  stud  for  years  at  a  time.  Not  every 
horse  may  be  affected.  It  has  been  connected  with  damp- 
ness, bad  fodder,  overwork  and  the  like.  Again  others 
have  connected  it  with  malaria  or  rheumatism. 
Potapenke,  Vigezzi,  Koch  and  others  have  found  various 
microorganisms,  no  two  of  which  seem  to  be  the  same. 
Even  an  animal  organism  like  malaria  has  been  described. 
(Whether  or  not  malaria  has  anything  to  do  w^th  the 
disease,  it  must  be  said  that  our  horse  was  favorably 
affected  in  regard  to  temperature  as  well  as  to  the  eye 
condition  by  repeated  subcutaneous  injections  of  Quinine 
Bisulphate,  Grain  xx  daily.)  The  attacks  last  five  to  nine 
days.  One  or  both  eyes  may  be  attacked  and  not  uncom- 
monl)^  do  they  alternate.  One  eye  may  cease  to  have 
attacks  while  the  other  continues.  The  experiments  here 
recorded  were  made  with  the  idea  of  transmitting  the 
disease  to  other  horses.  They  were  only  partially  suc- 
cessful. During  eight  months  the  affected  animal  referred 
to  us  had  six  attacks  of  ophthalmia.  The  attack  was 
observed  for  study  on  the  first  occasion,  but  during  the 
second  his  anterior  chamber  was  entered  by  a  needle 
attached  to  a  syringe,  the  exudate  aspirated  and  injected 
into  the  eye  of  a  horse  with  apparently  healthy  eyes.  The 
history  of  this  second  horse  will  be  given  later.  The 
attacks  of  the  first  horse  ranged  from  six  to  twelve  days. 
Five  of  the  six  affected  the  left  eye  and  one  the  right.  In 
January,  1910,  the  left  eye  was  used  for  further  inocu- 
lation, and  following  this  traumatism  complete  recovery 
never  took  place.  The  corneal  scar  left  by  the  needle  tract 
almost  disappeared,  but  an  inferior  anterior  synechia 
formed  and  was  followed  -by  a  spreading  opacity  of  the 


THE  EYE  407 

cornea,  much  wrinkling  of  the  iris  and  opacity  of  the 
depths.  After  the  fourth  attack  in  this  eye  it  was  com- 
pletely blind.  Material  was  obtained  from  this  eye  during 
its  last  attack,  but  it  was  merely  serous  fluid  containing 
a  few  blood  cells  and  epithelium,  but  no  bacteria. 

*'In  transferring  the  affection  from  this  animal,  the 
conjuctival  sac  was  washed  with  1-5000  bichloride  of 
mercury  solution  and  well  rinsed  with  salt  solution.  The 
anterior  chamber  was  then  entered  with  an  aspirating 
needle  and  the  exudate  removed.  This  consisted  of  0.4  cc. 
slightly  turbid  straw  colored  fluid  containing  a  few 
shreds  of  lymph.  Bacteriological  cultures,  moist  and  dry 
preparations  were  made  from  a  part  of  this,  while  the 
remainder  was  introduced  into  the  anterior  chamber  of 
the  second  horse.  This  animal's  eye  showed  the  effects 
of  the  traumatism  for  eight  days,  and  then  was  normal 
save  for  a  small  opaque  spot  in  the  cornea  left  from 
needle  puncture.  After  twenty-three  days  a  small  patch 
of  lymph  collected  in  the  pupil.  Tliis  increased  slowly 
accompanied  by  lacrymation  until  the  twenty-seventh 
day,  when  a  sudden  and  violent  conjunctivitis  arose.  The 
lymph  in  the  anterior  chamber  likewise  suddenly 
increased  and  rapidly  became  pus,  forming  a  hypopyon. 
The  conjunctivitis  became  purulent.  The  violent  stage 
lasted  five  days  and  slowly  subsided,  leaving  an  ectropion 
with  a  densely  injected  bulbar  conjunctiva,  almost  com- 
plete corneal  opacity  and  an  irregular  contraction  of  iris, 
apparently  due  to  several  small  synechias.  The  depths 
could  not  be  seen  because  of  the  corneal  condition.  This 
stage  of  affairs  remained  during  the  rest  of  the  animal's 
life,  two  months.  He  was  permitted  to  live  to  see  if  an 
exacerbation  of  this  chronic  process  or  involvement  of 
the  other  eye  would  appear.  Such  not  occurring  in  two 
months,  he  was  killed  and  the  eyes  removed.  Fluid 
removed  from  the  left  eye  of  the  first  horse  when  killed 
during  the  last  attack  was  injected  into  the  anterior 
chamber  of  a  third  horse.     This  animal's  eye  received 


408   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

the  operation  well  and  the  trauma  had  entirely  disap- 
peared when  the  animal  died  on  the  eighth  day. 

''Laboratory  Examinations. — From  fluid  removed 
from  horse  eye  (No.  1)  anaerobic  cultures  made  on  milk 
and  blood  serum,  blood  agar,  glycerine  agar ;  cultures  were 
made  directly  from  the  fluid,  while  the  coagula  were  dried 
upon  slides  and  stained  as  follows:  Loefifler's,  Gram's, 
Giemsa.  In  all  there  are  very  few  recognizable  bodies. 
They  are  red  blood  cells,  polynuclears  and  a  very  few 
small  mononuclear  cells.  In  regard  to  microorganisms 
three  structures  present  themselves.  A  well  staining 
Gram-positive,  rounded  end  rod  of  fairly  uniform  size  but 
tending  to  grow  in  pairs  and  stain  rather  irregularly  with 
Loeffler  and  Giemsa.  These  forms  are  sometimes  called 
'  dumbbell  '  in  that  they  are  bipolar,  or  even  seem  to  have 
a  constriction  in  their  centre.  Another  form  is  peculiar 
and  cannot  be  said  to  be  recognized  as  a  bacterium.  It  is 
circular,  of  fairly  regular  size  and  contour  and  in  many 
places  looks  like  a  very  large  coccus.  In  Loeffler 's  stain 
it  is  colored  deeply  in  the  centre  with  a  paler  marginal 
zone  and  an  unstained  halo  about  it,  which,  however,  is 
not  like  a  capsule.  In  the  Gram  and  Giemsa  method  it  is 
deeply  blue  or  purple  with  a  refractile  centre  and  very 
sharply  outlined  contour.  These  forms  varied  from  3 
to  5  microns.  The  third  form  is  a  wavy  delicate  short 
mycelium-like  thread.  Smears  from  the  cultures  as  made 
above  showed  chiefly  a  Gram-positive,  rounded  end  rod 
but  which  did  not  grow  on  planting  out.  It  grew  on 
aerobic  media,  but  was  not  found  on  anaerobic.  The 
Gram-positive  organism  would  not  grow  beyond  the 
fourth  generation.  It  was  not  identified  with  any  known 
species  by  the  characters  manifested  during  the  short 
time  we  were  able  to  keep  it  alive  but  could  be  placed  in 
the  Hog  Cholera  group.  The  mycelium  was  found  to  be 
an  aspergillus.  In  regard  to  the  large  coccus-like  body, 
little  can  be  added  to  the  above  description.  Further 
examination  did  not  reveal  characters  permitting  us  to 


THE  EYE  409 

place  it  among  the  protozoa.  No  evidences  of  division 
were  seen.  The  body  is  quite  uniform  in  appearance, 
varying  only  in  size.  Whatever  this  is  it  seems  to  be  an 
organized  body. 

''Cultures  from  pus  in  the  anterior  chamber  of  the 
second  horse  showed  the  palely  staining  rod,  an  asper- 
gillus  and  Micrococcus  aquatilis.  The  first  was  planted  on 
horse  serum  bouillon,  but  did  not  grow  after  the  first  gen- 
eration. These  cultures  were  made  after  death,  but  the 
cultures  made  during  the  acute  attack  direct  from  con- 
junctival sac  contained  such  a  host  of  organisms  that  no 
judgment  could  be  formed  of  their  relative  importance. 
The  polar  staining  rod  was  found  in  smears.  No  large 
coccus-like  bodies  were  observed  in  the  second  horse. 
Fluid  taken  from  the  first  horse 's  eye  at  death  was  sterile. 

''These  observations  are  at  variance  with  those  of 
others  but  such  results  are  not  unique  in  this  respect.  It 
seems  as  if  the  polar  staining  rod  deserves  some  consid- 
eration, and  we  expect  to  devote  some  attention  to  it  if 
another  horse  suffering  from  recurrent  ophthalmia  come 
to  our  notice.  The  large  coccus-like  bodies  are  very  inter- 
esting and  may  be  protozoa.  The  finding  of  the  amoeba 
in  the  cases  of  Potapenke,  increases  their  importance. 
Before,  during  and  after  the  fourth  attack  of  the  first 
horse  twenty  grains  of  quinine  bisulphate  were  given 
hypodermically  daily  for  twenty  days.  The  attack  was 
very  mild.  Before  the  drug  was  given  his  temperature 
had  ranged  from  99°  to  101°  F.  Immediately  after  the 
first  dose  the  temperature  fell  to  below  99°  F.,  and 
remained  at  a  very  regular  level  during  the  entire  twenty 
days.    No  malarial  organisms  were  found  in  the  blood. ' ' 

The  ear  is  without  special  interest  except  as  a  place 
of  localization  of  sarcoptes,  demodex  and  fly  larvae.  A 
few  cases  of  acute  catarrhal  otitis  media  have  been  found 
in  association  with  nasopharyngitis  both  of  the  non- 
specific variety  and  that  which  resembles  distemper.  One 
case  which  led  to  meningitis  has  been  mentioned. 

27 


SECTION  XIV 
CONSTITUTIONAL  DISEASES 

Theee  is  a  long  list  of  diseases  including  among  others 
such  conditions  as  hyperthyroidism,  osteodytrophies, 
diabetes  and  gout  which  are  spoken  of  as  constitutional 
but  which  in  reality  are  usually  dependent  upon  some 
lesion  peculiar  to  a  definite  organ.  Several  have  been 
discussed  under  systemic  diseases  so  that  there  remain 
for  consideration  in  this  section  only  two,  gout 
and  diabetes. 

Constitutional  diseases  are  recognized  in  mid  animals 
either  not  at  all  or  by  some  happy  chance  which  permits  of 
examination  direct  enough  to  elicit  diagnostic  criteria. 
Gout  has  been  discovered  for  example  in  some  parrots 
and  herons  because  of  their  swollen  feet  and  their  move- 
ments. In  veterinary  practice  fairly  accurate  diagnoses 
are  possible  but  in  wild  collections  they  are  nearly  always 
hit  or  miss.     Therapeutics  naturally  follow  this  rule. 

Gout. 

Gout  in  mammals  has  been  observed  in  the  London 
Zoological  Garden  but  has  not  been  encountered  here  or 
we  have  overlooked  it.  A\aan  gout  on  the  other  hand  in 
one  of  its  forms  comes  to  our  attention  not  infrequently. 
;'It  occurs  most  often  in  parrots,  gallinaceous  and  anserine 
birds  and  herons ;  occasionally  accipitrine  birds  will  suffer 
with  it,  an  observation  more  often  recorded  in  European 
collections  than  with  us.  The  figures  show  no  pre- 
dominance of  percentage  for  any  order  but  the  records 
indicate  that  the  most  beautiful  examples  of  internal 
uratic  deposits  occur  in  the  anserine  birds  and  parrots, 
while  the  best  specimen  of  general  gout,  including  the 
joints,  was  found  in  a  boat-billed  heron  {Cancroma 
cochlearia)  quoted  below.  1 

410 


CONSTITUTIONAL  DISEASES  411 


In  so  far  as  etiology  of  this  disease  is  concerned  in 
domestic  stock,  too  rich  food,  especially  in  protein,  and 
restriction  of  activity  seem  to  be  credited  with  the 
greatest  influence.  These  factors,  while  doubtless  of 
importance  for  birds  as  they  are  believed  to  be  for  man, 
do  not  seem  to  fill  all  the  requirements  since  all  our 
specimens  are  confined  and,  because  of  their  lack  of  exer- 
cise, possibly  receive  too  much  food,  /judging  by  our 
observations  and  by  publications  from  other  gardens, 
carnivorous  birds  are  not  conspicuous  for  the  incidence 
of  gout  whereasj/grain-  seed-  and  fish-eaters  suffer  more 
often.  /This  suggests  that  these  varieties  cannot  dispose 
of  dierary  protein  which  might  be  excessive  for  their 
metabolism  while  in  captivity,  whereas  carnivorous 
species  have  a  digestive  and  chemical  reserve  to  take  care 
of  excess  protein.  Some  such  accommodative  power  must 
exist  in  human  beings  since  not  every  large  meat-eater 
develops  gout.  Heredity,  often  blamed  for  the  human 
disease  cannot  help  us  with  these  birds.  Examination  of 
the  diet  list  at  the  Garden  does  not  reveal  a  great  per- 
centage of  concentrated  protein  in  the  feed  of  the  grain- 
and  seed-eaters.  The  disease  occurs  too  seldom  to 
disturb  the  accepted  dietary  for  its  possible  elimination. 
Studies  now  going  on  may  indicate  appropriate  changes 
in  the  dietaries  that  might  be  responsible. 

Arthritic  gout  appears  usually  in  the  pedal  joints 
but  may  be  found  in  the  wings.  Irregular,  sometimes 
very  deforming  swellings  appear  which  must  be  tender 
judging  by  the  quietness  of  the  bird  and  by  its  behavior 
if  the  joints  be  touched,  "i  Most  often  the  swelling  seems 
greater  upon  the  flexor  (palmar)  surfaces  of  the  toes  or 
in  the  end  of  the  tarsal  articulation.  Aside  from  these 
few  observations  there  is  notliing  peculiar  about  the  at- 
tack or  the  specimen  during  its  sickness.  Chronicity  seems 
to  be  the  rule  and  little  emaciation  may  be  found. 
Appetite  is  normal  or  excessive,  provided  the  food  can 
be  reached. 


412   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Internal  or  serous  membrane  gout  cannot  be  recog- 
nized during  life  so  far  as  I  know.  The  bird  may  seem  in 
its  usual  condition  of  feather,  activity,  appetite  and 
elimination,  when  suddenly  it  will  fade  in  a  day  or  so  and 
die.  At  autopsy  the  serous  surfaces  of  the  heart  and 
peritoneum  will  be  white  with  uric  acid  crystals  and  the 
kidneys  a  pale  yellow  brown  with,  markings  indicating  that 
the  pelves  and  tubules  are  choked  with  urates. 

The  boat-billed  heron  (Cancroma  cochlearia)  had  had  bad  feet 
for  three  months.  The  general  condition  is  poor  as  to  plumage  and 
flesh.  The  tarsal  and  metatarsal  joint  areas  of  both  legs  are  sur- 
rounded by  firm  tough  swellings  involving  skin  and  periarticular  tissue. 
That  on  left  foot  has  ulcerated  and  bled.  On  section  the  swelling  is 
found  to  consist  of  reddened  fibrous  tissue  around  tendons,  the  latter 
apparently  running  through  smooth  sheaths.  At  both  ankles  are  urate 
deposits  clearly  seen  in  this  inflammatory  tissue  but  at  the  lower  end 
of  the  tarsus  there  are  no  distinct  deposits.  The  joint  surfaces  do  not 
seem  to  be  involved.  Knee,  hip,  and  wing  joints  seem  uninvolved. 
Internally  all  surfaces  are  opaque  by  sprinkling  of  whitish  or  yellowish 
dots  like  urates ;  this  is  especially  marked  over  heart.  Pleurae  aside  from 
urates  are  negative.  Lungs  very  slightly  uniformly  congested  through- 
out. Aorta  and  branches  are  stiff,  intima  smooth.  The  liver  is  soft, 
deep  brown  color,  architecture  seems  normal.  The  kidney  has  a  smooth 
capsule  and  a  smooth  pale  yellow  surface.  Organ  is  firm.  Section  sur- 
face is  glistening  and  opaque,  every  lobule  clear,  pelves  filled  with 
pale  yellow  material,  cortical  areas  irregular.  Alimentary  tract  nega- 
tive. Microscopical  section  of  kidney  shows  general  topography  re- 
tained, vessels  very  much  injected,  some  showing  thrombosis.  Cortex 
slightly  irregular  probably  by  swelling  of  medulla.  Tubular  epithelium 
swollen  and  granular  or  desquamating  and  degenerating.  Glomeruli 
vary  in  size  and  shape,  mostly  fill  out  the  capsule.  Capillary  cells  show 
some  vacuoles.  Some  urate  collections  in  tubules;  practically  all  pelvic 
tubules  have  some  urates.  Interstitial  tissue  not  increased.  Blood 
vessel  walls  somewhat  loose.  Endothelium  prominent.  No  areas  of 
degeneration  seen. 

Diabetes. 

Diabetes  is  an  infrequent  but  well  recognized  disease 
among  domestic  animals.  Its  detection  depends  on  a 
rather  vague  chain  of  symptoms  confirmed  by  the  dis- 
covery of  sugar  in  the  urine.  For  the  suspicion  that  a 
wild  animal  was  suffering  with  diabetes  one  would  have 


CONSTITUTIONAL  DISEASES  413 

to  rely  upon  great  thirst,  loss  of  flesh,  depression,  exces- 
sive urination  and  possibly  cataractous  opacity  of  the 
eye.  Such  a  chain  of  symptoms  has  not  been  detected. 
At  every  occasion  at  postmortem  that  the  bladder  is  full 
of  urine,  a  routine  examination  is  made.  In  this  way  we 
detected  one  case  which  seems  to  have  been  diabetes,  the 
diagnosis  being  based  upon  the  glucosuria  and  the 
lipemia.  For  some  unknown  reason  a  section  of  the  pan- 
creas was  not  made,  a  regrettable  matter  since  a  definite 
purulent  gingivitis  existed  and  may  have  lain  at  the  basis 
of  an  infective  pancreatitis,  well  known  to  be  the  cause 
of  certain  cases  of  diabetes.  The  case  is  recorded  in  full 
since  it  is  unique,  no  other  case  in  a  wild  animal  being 
fully  reported. 

The  arctic  fox  [Canis  lagopus)  ate  and  appeared  well  the  day 
before  it  was  found  dead.  Diagnosis — Diabetes  mellitus.  The  animal 
was  in  good  condition.  The  left  conjunctiva  was  reddened,  congested, 
edematous,  with  slight  mucopurulent  discharge  in  canthus.  Muscles 
have  a  cloudy  appearance.  Fat  lacks  rich  yellow  color.  The  general 
impression  of  anemia  is  present.  Lungs  and  pleura  are  normal  through- 
out. Heart  muscle  is  pale,  firm  and  tough.  The  tricuspid  shows 
thickening  of  the  edge  of  posterior  leaflet,  the  mitral  shows  slight  sclero- 
sis of  edge  of  mesial  leaflet.  The  auricles  are  distended  with  clot. 
Left  ventricular  wall  is  greatly  thickened.  Upon  incising  the  heart 
the  surface  of  blood  shows  fine  fat  globules.  Peritoneum  is  normal. 
Liver  is  slightly  increased,  surface  smooth,  edges  rounded,  consistency 
soft,  color  brownish  red  with  yellow  mottlings  which  are  without  defi- 
nite boundaries ;  the  section  surface  is  moist,  granular  and  opaque.  The 
bile  is  fluid,  green-yellow  and  the  duct  is  patulous.  The  spleen  is 
slightly  enlarged  and  soft.  The  kidney  is  slightly  enlarged,  capsule 
strips  easily  leaving  a  smooth,  purplish  red  surface;  section  surface 
is  glistening,  moist  and  exudes  blood;  consistency  is  slightly  softened; 
cortical  striae  very  distinct.  The  bulging  cut  surface  and  poor  demarka- 
tion  of  cortex  and  medulla  characteristic  of  acute  nephritis  are  present. 
The  organ  shows  fat  globules  in  the  expressed  blood.  The  adrenals 
are  very  small,  firm,  brown,  bean-shaped  bodies  with  a  brownish  medulla. 
The  bladder  is  slightly  distended  with  turbid  urine.  Urine  shows  dark 
granular  casts,  compound  granule  cells,  spermatozoa  and  a  positive 
Fehling's  test.  Prostate  is  large  and  firm  and  a  turbid  material  exudes 
from  external  meatus.  The  mouth  shows  several  decayed  teeth.  In 
the  neighborhood  of  last  molars  on  left  side  of  upper  jaw  a  bead  of  pus 
exudes;  further  pressure  results  in  no  greater  flow.     The  stomach  is 


414  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

distended  with  a  great  quantity  of  undigested  food  and  gas ;  no  worms. 
Serosa  and  wall  normal  but  anemic.  Duodenum  normal.  Jejunum  con- 
tains numerous  worms  about  1  to  1.5  cm.  long ;  it  is  distended  with  gas. 
The  pancreas  is  large,  soft,  like  fat,  white ;  it  extends  between  the  layers 
of  mesentery  along  the  course  of  the  duodenum;  at  first  the  pancreas 
was  mistaken  for  fat. 

Histological  Notes. — Spleen  shows  a  distinct  overgrowth  of  trabec- 
ulaB.  Beyond  this  there  is  nothing  pathological.  Liver  shows  distended 
portal  venules  in  which  there  are  chains  of  bacilli.  There  is  no  especial 
fibrous  overgrowth  of  capsule  of  Glisson;  capillaries  are  choked 
with  shadow  corpuscles  and  here  too,  long  chains  of  bacilli  may  be  seen ; 
parenchyma  cells  show  postmortem  change.  Adrenal  is  the  seat  of 
postmortem  degeneration,  not  congested,  nor  is  there  any  evidence  of 
bacterial  invasion.  The  kidney  shows  no  interstitial  changes,  in  fact 
the  section  seems  to  be  entirely  normal  save  for  moderate  congestion. 
Vessels  show  no  bacteria. 


SECTION  XV 
THE  RELATION  OF  DIET  TO  DISEASE 

BY 

Db.  E.  p.  CoRSOISr-WHITE 

Food  in  the  widest  acceptation  of  the  term,  means 
every  thing  ingested  that  goes,  directly  or  indirectly,  to 
growth,  repair  of  the  body,  or  production  of  energy,  all  of 
which  phenomena  must  continue  when  food  is  mthheld  or 
supplied  in  insufficient  quantities.  Under  the  latter  con- 
dition the  processes  go  on  at  the  expense  of  the  body 
tissues  as  these  are  protected  only  when  the  diet  is  ade- 
quate in  every  way.  A  proper  diet,  therefore,  must  be 
one  on  which  an  animal  will  attain  maximum  development, 
maintain  a  normal  weight  curve,  show  a  minimum  sus- 
ceptibility to  disease,  live  out  a  full  term  of  life,  breed 
normally,  and  rear  healthy  offspring,  capable  of  normal 
independent  life  after  they  are  weaned.  It  must  fulfiU 
the  caloric  needs  of  the  body,  and  in  young  animals  it  must 
also  supply  the  growth  impulse.  In  its  physical  prop- 
erties it  must  fit  the  morphological  demands  of  each  type 
of  gastrointestinal  tract.  In  its  chemical  content  it  must 
supply  all  the  elements  found  in  the  body  in  usable  form, 
and  in  amounts  sufficient  to  cover  the  needs  of  the  body 
for  growth,  repair  and  waste.'  To  evaluate  fully  the 
influence  of  food  on  the  individual  animal  it  is  necessary 
to  study  its  relation:  (1)  to  the  type  of  alimentary  tract, 
(2)  to  the  type  of  bacterial  flora  and  their  metabolic 
processes,  (3)  to  the  chemical  needs  of  the  body,  (4)  to 
the  changes  arising  in  the  catabolism  and  anabolism  of 
all  types  of  food,  (5)  to  exercise  or  its  lack,  keeping  in 
mind  always  the  constant  interdependence  of  all  factors. 
Our  knowledge  of  nutrition  has  to  a  very  large  extent 

415 


416   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

paralleled  the  advances  in  chemistry,  especially  the  re- 
searches into  the  structural  make-up  of  living  cells,  the 
intermediate  stages  in  their  upbuilding  and  degradation 
and  the  products  resulting  from  their  physiologi- 
cal activities. 

Incorrect  feeding  both  qualitative  and  quantitative  un- 
doubtedly plays  an  important  role  in  producing  disease. 
In  the  early  works  on  nutrition,  the  proportion  of  fats, 
carbohydrates  and  proteins  was  regarded  as  the  essential 
point  of  a  normal  diet.  The  researches  on  the  composi- 
tion of  foods  marked  the  first  real  epoch  in  this  .history 
and  Fischer's  (1)  studies  on  the  variation  in  the  composi- 
tion of  proteins  from  different  sources  first  introduced 
the  idea  of  quality.  Later  Mendel  and  Osborne  investi- 
gated the  biological  values  of  purified  proteins,  while  at 
the  same  time  McCollum  and  others  were  studying  the 
value  of  the  groups  of  proteins  occurring  in  a  single 
natural  food  stuff,  were  calling  attention  to  the  so-called 
vitamines,  and  were  emphasizing  the  need  of  balanced 
inorganic  materials.  These  studies  have  practically 
revolutionized  our  knowledge,  particularly  of  the  effects 
of  badly  balanced  foods.  They  have  clearly  demonstrated 
that  dietary  values  can,  in  all  probability,  be  discovered 
only  by  careful  biological  study  of  feeding  experiments 
together  with  the  finer  analysis  of  the  components  of  the 
diet,  especially  of  the  protein  and  fat  radicles.  At  the 
same  time  a  definite  appreciation  of  the  role  of  each  ele- 
ment in  metabolism  must  be  kept  in  mind. 

These  varied  studies  on  nutrition  have  shown  that  the 
chemical  requirements  of  a  diet  are  in  their  ultimate 
analysis  essentially  the  same  for  all  species  of  the  higher 
animals — that  is  all  require  approximately  the  same 
amount  of  protein,  fat,  carbohydrate,  etc.,  per  kilo  of  body 
weight,  while  the  morphology  of  the  tract  decides  the 
physical  properties  of  the  diet. 

(1)    Chemistry  of  the  Proteins,  Mann. 


THE  RELATION  OF  DIET  TO  DISEASE       417 

Relation  of  Food  to  Alimentary  Tract. 

Food  derived  from  animal  sources  is  high  in  protein, 
readily  digested,  and  highly  putrefactive.  This  type  of 
diet  is  suited  to  an  alimentary  tract  which  permits  rapid 
passage  through  its  leng-th,  and  is  fitted  with  sturdy  walls. 
The  gastric  section  is  simple,  the  intestine  short  and 
narrow  with  ill-defined  separation  of  its:  parts  into  small 
gut,  cecum  and  colon.  This  type  is  found  in  all  land 
Carnivora.  The  fish-eating  carnivores  have  a  strong 
tubular  stomach  and  an  enormous  length  of  intestine,  but 
no  cecum.  The  omnivores  occupy  a  middle  place.  In 
them  the  alimentary  tract  consists  of  a  simple  stomach,  a 
short  wide  intestinal  tube,  and  a  more  complex,  although 
still  comparatively  simple,  cecum  which  is  generally 
longer  than  that  found  in  the  carnivores.  This  tract  is 
too  small  to  manipulate  the  bulky  vegetable  masses  neces- 
sary to  pro\'ide  their  minimum  protein  requirement,  and 
too  long  and  complicated  to  dispose  quickly  of  the  putre- 
factive animal  tissue.  Among  these  animals  colitis  is 
common,  due  to  the  fact  that  the  shape  and  position  of  this 
part  of  the  tract  favors  stasis,  or  at  least  a  sluggish 
movement  of  its  contents  at  a  point  in  the  digestive 
scheme  where  the  food  residue  is  rich  in  protein  by- 
products, ready  for  bacterial  gro^vth. 

The  herbivores  mth  food  derived  from  plants  which 
requires  a  long  period  of  time  for  its  digestion,  have,  on 
the  other  hand,  voluminous  stomachs,  or  large  ceca  or 
both;  and  very  long  small  intestine.  In  this  tract  the 
concentrated  food  of  the  carnivores  would  provide  an 
enormously  excessive  protein  intake  or  if  only  the  protein 
requirement  is  supplied  would  leave  the  tract  so  empty 
that  it  would  be  unable  to  functionate. 

All  studies  in  comparative  anatomy  demonstrate  the 
fact  that  while  neither  a  complex  stomach  nor  a  large 
cecum  is  essential  to  the  digestion  of  vegetable  food,  a 
capacious  and  complex  alimentary  canal,  as  a  whole,  bears 
a  relation  to  vegetable  diet,  particularly  in  the  mammals. 


418  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Either  a  highly  developed  concentrated  glandular  appa- 
ratus is  added  to  the  stomach,  as  in  the  wombats,  beavers 
and  dormice,  or  the  stomach  is  subdivided,  sacculated,  or 
otherwise  amplified  as  in  the  ruminants  and  herbivorous 
marsupials.  Sometimes  both  complexities  are  combined 
as  in  the  case  of  the  sloths.  If  the  simple  stomach  is 
retained,  it  is  supplemented  by  a  large  sacculated  colon  or 
cecum,  as  in  the  horse.  In  birds,  the  proventricle  is  larger 
in  meat-  and  fish-eaters,  while  the  gizzard  is  more  muscu- 
lar in  grain-  and  insect-feeders,  and  the  intestines  are 
longer  in  those  devouring  coarse  green  grass  and  leaves. 
The  length  of  the  ceca  is  related  entirely  to  the  diet,  the 
long  ones  corresponding  to  the  diet  which  needs  pro- 
tracted periods  of  time  to  exhaust  its  nutriment,  j 

The  Bacterial  Flora. 

'  The  bacterial  flora  harbored  in  the  intestinal  tract  is 
closely  related  to  the  type  of  food  and  to  the  character  of 
the  alimentary  tract.  Levin(2)  found  sterile  intestinal 
tracts  in  white  bears,  seals,  reindeer,  eider  ducks  and 
penguins  when  in  the  Arctic  regions;  but  these  same 
animals  when  they  are  brought  to  a  temperate  climate 
rapidly  acquire  intestinal  bacteria.  The  function  of  the 
normal  inhabitants  of  the  tract  is,  probably,  to  protect 
the  body  against  invasions  of  obnoxious  species.  Herter 
found  in  man  that  a  few  species  adapt  themselves  to  the 
digestive  tract  and  control  the  growth  of  new-comers 
capable  of  doing  injury.  These  common  varieties  become 
a  source  of  danger  only  when  present  in  large  numbers. 

Bacteria  which  produce  decomposition  of  food  in  the 
digestive  tract  are  of  three  types  :  (1)  Pure  putrefactive 
anaerobes,  (2)  organisms  both  fermentative  and  putre- 
factive, but  tending  generally  to  antagonize  the  putrefac- 
tive anaerobes,  and  (3)  fermentative  organisms.  In  the 
stomach,     fermentation     of     carbohydrates     vnth.     the 

(2)  Ann.  Inst.  Past.,  1S99,  XIII,  558,  and  Skandinavisches  Arch.  f. 
Physiol.,  1904,  XVI,  249. 


THE  RELATION  OF  DIET  TO  DISEASE       419 

production  of  organic  acids  is  a  frequent  occurrence.!; 
Putrefactive  types  are  very  rare  except  with  pyloric 
stenosis,  a  condition  which  favors  excessive  fermentation 
by  diminishing  the  tone  and  motility  of  the  stomach  and 
the  amount  of  hydrochloric  acid.  This  condition  is 
further  increased  by  excessive  carbohydrate  food.  In 
general  the  products  of  fermentation  tend  to  restrict 
putrefaction,  yet  both  may  be  operative.  In  the  small 
intestines,  bacteria  are  always  present  because  of  the  pro- 
tein richness  of  secretions,  the  rapid  digestion  of  food  and 
the  slight  or  ineffectual  antiseptic  properties  of  intestinal 
juice,  bile  and  pancreatic  secretions.  The  putrefactive 
bacteria  rapidly  increase  and  decompose  any  protein  that 
is  miabsorbed — a  process  most  marked  in  the  colon 
because  its  shape  and  position  favor  stasis  or  slow  move- 
ment of  its  contents.  In  general  the  greater  the  amount 
of  unabsorbed  and  digestible  protein  and  the  longer  the 
material  stays  in  the  intestinal  tract,  the  greater  the 
putrefaction.  The  meat-eating  animals  develop  Gram- 
negative  bacilli,  while  the  carbohydrate-eaters  show  a 
predominance  of  Gram-positive  types. 

Ingested  food  never  contains  the  enormous  amount  of 
bacteria  found  in  the  feces.  The  alunentary  tract  with  its 
contents  forms  a  most  efficiently  combined  incubator  and 
culture  medium,  in  which  bacterial  growth  exceeds  that  of 
any  known  location  both  in  intensity  and  complexity.  The 
range  of  reaction  and  composition  of  nutritive  substances 
at  different  levels  of  the  intestinal  tract  is  such  that  a 
great  variety  of  bacteria  capable  of  growth  at  body  tem- 
perature develop.  The  prominent  types  that  appear  in 
the  flora  of  each  order  of  mammals  are  fairly  constant 
in  their  occurrence.  They  depend  primarily  on  food 
ingested,  and  show  well  marked  seasonal  variations, 
dependent  again  on  changes  in  food.  Faulty  feeding  may 
itself  give  rise  to  a  toxic  condition  of  the  gastrointestinal 
tube,  and  thus  often  prepares  this  soil  for  the  develop- 
ment of  organisms. . 


420   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

The  intestinal  flora  also  changes  along  rather  definite 
lines  as  the  diet  of  the  host  changes  from  the  monotony  of 
the  infant  to  the  variety  of  the  adult.  At  birth  the  tract 
is  sterile,  but  bacteria  soon  make  their  entry  through 
the  mouth  in  food  and  water.  The  majority  of  these 
organisms  pass  to  the  stomach  where  many  are  destroyed, 
but  a  number  travel  to  the  intestines  where  they  may  gain 
a  foothold.  There  is  always  a  mechanical  transportation 
of  intestinal  bacteria  from  higher  to  lower  levels.  A  con- 
tinued preponderance  of  protein  in  the  diet  of  all  animals 
leads  to  a  partial  or  complete  suppression  of  the  Gram- 
positive  acid-forming  groups  and  an  increase  of  the  pro- 
teolytic Gram-negative  types ;  while  on  the  other  hand  an 
excess  of  carbohydrate  leads  to  diminution  or  suppression 
of  proteolytic  activity  and  an  increase  in  the  fermentative 
organisms.  Therefore  the  most  important  normal  factor 
in  determining  the  intestinal  flora  in  health  is  the  chem- 
ical composition  of  the  ingested  foods. 

The  nature  of  the  dominant  organisms  which  develop 
in  diets  rich  in  carbohydrates  varies  with  the  carbo- 
hydrate itself.  In  all  ordinary  diets  there  are  (1)  starches 
— forms  not  readily  fermentable,  and  (2)  sugars — which 
are  largely  absorbed  from  the  higher  levels  of  the  small 
intestine,  leaving  residual  starches  and  proteins  in 
relatively  great  concentration  in  the  lower  levels.  There- 
fore the  obligate  fermentative  organisms  are  prominent 
only  in  the  higher  levels,  the  facultative  appear  in  the 
intermediate  places,  and  the  obligate  proteolytic  organ- 
isms in  the  lower  intestines.  This  accounts  in  a  measure 
for  the  great  increase  of  lower  intestinal  disturbances  in 
omnivores.  Complete  proteins  resist  putrefaction,  but 
the  products  of  protein  digestion  and  of  the  intestinal 
secretions  constitute  the  main  substrata  for  putrefactive 
bacteria.  Animal  protein  develops  more  active  proteo- 
lytic bacteria  than  vegetable  protein,  which  accounts  for 
the  greater  predominance  of  putrefactive  infections  in 


THE  RELATION  OF  DIET  TO  DISEASE       421 

There  are  two  important  factors  to  consider  in  dis- 
cussing the  influence  of  diet  on  intestinal  bacteria: 
(1)  The  substitution  of  types,  which  frequently  follows  a 
monotonous  diet,  and  (2)  the  change  in  metabolism  of 
existing  types  of  bacteria  when  dietary  conditions  are 
such  that  the  intestinal  medium  at  one  or  another  level 
fluctuates  in  its  content  of  usable  carbohydrate  and  other 
nutrient.  The  nature  and  extent  of  these  modifications 
and  their  effects  upon  the  host  vary  greatly,  not  only 
qualitatively  but  quantitatively.  An  invasion  of  the  tract 
by  exogenous  bacteria,  as  the  dysentery  bacillus,  cholera, 
typhoid,  etc.,  in  food  or  water  may  lead  to  a  more  or  less 
pronounced  replacement  of  some  of  the  normal  intestinal 
tjipes  by  these  alien  organisms,  and  to  the  production 
of  disease. 

The  importance  of  all  the  foregoing  facts  concerning 
the  changes  in  the  food,  in  the  intestinal  cultural  substrata 
and  in  the  advent  of  new  kinds  of  organisms  was  emphati- 
cally demonstrated  in  the  marked  fall  in  gastrointestinal 
diseases  in  carnivores  after  proper  screening  of  meats. 
The  simple  protection  of  the  food  given  to  these  animals 
eliminated  the  air  bacteria  which,  entering  from  dust  and 
flies,  alter  the  chemistry  of  the  meat  before  consumption 
or  change  the  flora  of  the  intestine  after  consumption. 
Normal  organisms,  or  types  indistinguishable  from  them, 
may  multiply,  through  unusual  conditions,  extend  their 
normal  habitat,  and  eventually  lead  to  abnormal  reactions 
detrimental  to  the  host.  These  facts  throw  considerable 
light  on  the  site  and  character  of  gastrointestinal  lesions 
found  in  various  orders,  a  subject  to  be  discussed  more 
fully  later. 

There  are  many  intestinal  disturbances  of  unknowni 
causation,  in  some  of  which  bacteria  presumably  play  a 
secondary  part.  The  primary  disturbance  is  due  to  the 
products  resulting  from  the  action  of  bacteria  upon  food. 
(Many  toxic  bodies  are  produced  either  before  or  after 
ingestion  by  the  bacterial  decomposition  of  carbohydrate. 


422   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

fat  or  protein,  independent  of  any  actual  infection.  The 
s^Tuptoms  arising  from  bacterial  decomposition  of  foods 
depend  largely  on  the  organism  concerned  and  vary  from 
a  mild  intoxication  to  a  severe  toxemia.; 

Relation  of  Dietaey  Groups  to  Autopsy  Diagnoses. 

Analysis  of  the  autopsies  on  file  from  sole  point  of 
view  of  dietary  habits  of  the  animals  gives  rather  inter- 
esting groupings  of  disease  states,  which  apparently  and, 
in  some  cases  definitely,  emphasize  the  relationship 
between  food,  metabolism  and  disease.     (Table  19.) 

From  this  table  a  few  facts  stand  out  prominently. 
It  is  definitely  sho^vn  that  both  birds  and  mammals  on  a 
diet  of  mixed  animal  and  plant  tissue  show  a  low  per- 
centage of  disease  in  the  gastrointestinal  tube,  liver, 
pancreas  and  kidney.  The  mammals  on  this  diet  give  the 
highest  figures  for  anemias  and  degenerative  osseous 
conditions.  Birds  on  this  diet  show  very  little  osteo- 
malacia, but  a  fair  amount  of  anemia.  Possibly  this  may 
be  accounted  for  by  the  fact  that  all  of  them  pick  gravel 
and  may  be  able  from  this  to  supply  some  of  the  inor- 
ganic deficiency.  Carnivorous  birds  and  mammals,  on  the 
other  hand,  show  an  exceedingly  large  assortment  of 
gastrointestinal  disorders,  diseases  of  the  accessory 
glands  of  digestion,  and  of  the  kidneys.  Disorders  of  the 
thyroid  gland  are  almost  entirelj^  confined  to  carnivorous 
mammals — 7.5  per  cent.,  compared  to  0.25  per  cent,  in  all 
other  orders.  Gout,  while  common  among  birds,  was  not 
present  in  any  mammalian  autopsy,  while  arthritis  in 
mammals  reached  its  highest  record  among  grass-  and 
grain-eating  herbivora.  The  percentage  of  rickets  was 
highest  in  the  young  carnivores  (2.6  carnivores  as  against 
.4  per  cent,  in  all  other  mammals),  and  was  very  rare 
among  all  birds. 

The  succulent  vegetable  diet  was  lowest  in  its  relation 
to  degenerative  visceral  disorders  and  highest  in  acute 
gastritis ;  the  latter  fact  was  probably  due  to  the  fermen- 


THE  RELATION  OF  DIET  TO  DISEASE       423 


tation  of  soft  moist  food  that  requires  rather  a  long  time 
for  its  primary  digestion.  This  type  of  food  has  also  a 
high  and  easily  available  sugar  content  which  makes  it  a 

Table  19. 
An  Analysis  of  the  Pathological  Findings  Described  in  the  5,365  Autopsies  from 
the  Point  of  View  only  of  the  Dietary  Habits  of  the  Animals.  The  Percentage 
Results  Represent  the  Proportionate  Number  of  Cases  of  Each  Pathological 
Lesion  Found  in  the  Entire  Group  of  Animals  on  Each  Special  Diet  without 
Reference  to  Zoological  Orders.  


Disease  states 


Malnutrition 

Food  Poisoning 

Acute  Gastritis 

Acute  Duodenitis .  .  .  . 

Acute  Enteritis 

Acute  Gastroenteritis . 

Chronic  Gastritis 

Chronic  Enteritis .  .  .  . 

Colitis 

Acute  Pancreatitis.. .  . 
Chronic  Pancreatitis. . 
Acute  Liver  Disease. . . 
Chronic  Liver  Disease 

Acute  Nephritis 

Chronic  Nephritis. .  .  . 
Myocardial     Degene- 
ration   

Arterial  Disease 

Anemia  pernicious..  .  . 
Anemia  secondary. . . . 

Thyroid  Disease 

Adrenal  Disease 

Diabetes 

Osteomalacia 

Osteitis  deformans. . .  . 

Arthritis 

Rickets 

Gout 

Sore  Eyes 

MaUgnancy 

Tuberculosis 


Mammalia  1860 


.1 
.3 

3.2 
.5 

2.5 
26.3 

1.1 

2. 

1.9 
.1 
.5 
.8 

3. 

9.1 

4.5 

.1 
.1 
.3 

4.2 

1.6 
5.2 


.05 
32.6 


6.3 
.3 
3.4 
53.2 
6. 
5.6 

2.2 
1.7 
1.3 
6.3 
12.2 
11.6 

.34 
3.1 
.32 
.32 
7.5 
1.3 
.2 
.4 


.3 

.9 

3.5 


9.3 

3. 

19.9 
2. 
3. 

1. 

.3 
3.3 

12.7 
6.7 


2.2 
2.5 
3.1 

.5 

3.1 

29.2 

.8 
2.2 

3.1 

1.4 

6. 

12.4 

7.8 

1.1 
2.2 

1.5 

.7 
1.5 


.05 
.05 
.9 
.1 

7. 
25.3 
.2 

1.1 

.4 
.2 

4.2 

1.1 

5.1 

2.9 

.3 
.3 

1.1 
.3 


12. 


.4 
.2 

2. 

1.4 

1. 
38.6 

1.4 

3.3 


1.2 
2.8 
2.5 
6.7 
6.7 

2. 
3.1 

2.5 
.2 


1.3 
1.2 
8. 
35.6 
.3 
1.3 


.5 
2.5 
1.6 
4.1 
2.1 


2.8 


17.2 


13.5 
5.4 
5.4 

64.8 
5.4 

13.5 


2.7 
13.5 

8.1 
13.5 

8.1 
1.8 


5.7 


very  favorable  medium  for  many  of  the  fermentative 
types  of  bacteria.  Most  of  the  lesions  in  this  group  were 
around  the  pylorus  and  upper  duodenum. 


424   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Overeating  is  a  factor  that  must  be  borne  in  mind  when 
considering  the  hay-  and  grass-eating  herbivora.  Packing 
of  the  rumen  is  a  not  infrequent  discover5\  This  condi- 
tion is  also  found  in  certain  seed-eating  birds.  As  a 
supply  of  food  is  constantly  at  the  disposal  of  these  ani- 
mals and  exercise  is  prevented  by  captivity,  continuous 
eating  becomes  their  principal  diversion.  In  this  group 
also  food  poisoning  was  highest,  a  condition  which  may 
be  due  to  (1)  spoiled  food,  (2)  poisonous  substances  in  the 
foods,  (3)  fermentation  of  grass  foods  (spoiled  hay  or 
musty  fodder).  Malnutrition  also,  is  higher  than  with  any 
other  diet,  due  probably  to  the  somewhat  meagre  nutri- 
tious value  of  the  food.  This  group  also  shows  a  high  per- 
centage of  acute  pancreatitis,  degeneration  of  the  liver, 
myocardimn  and  arteries.  Arthritis  was  present  in  this 
group  2.2  per  cent.,  against  0.2  per  cent,  in  all 
other  groups. 

A  study  of  Table  19  demands  a  constant  recollection 
of  the  morphology  of  the  tract  involved  and  its  main 
points  of  vulnerability,  the  bacteria  capable  of  living  on 
the  particular  type  of  food  or  its  constituents  and  the 
by-products  produced  during  the  digestion  and  absorp- 
tion of  these  foods.  Not  one  of  these  factors  can  be 
ignored  in  evaluating  the  influence  of  diet,  which  to  be 
correct  must  supply  elements  in  proportions  that  are 
chemically  available  for  body  needs  (for  instance.  Von 
Wendt(3)  found  that  more  iron  was  required  if  the  diet 
was  deficient  in  calcium).  These  proportions  must  be 
worked  out  by  carefully  combined  chemical  and  biologi- 
cal experiments. 

Malnutrition. 

There  was  one  omnivorous  beast,  a  Hamadryas 
Baboon,  which  represented  the  only  true  case  of  starva- 
tion, probably  induced  by  nostalgia,  as  it  never  ate  after 
coming  into  the  Garden.  Thirty  cases  of  partial  starva- 
tion   or    malnutrition    are    listed    in    our    records,    the 

(3)    Skandinavisches  Arch,  fiir  Physiologie,  Vol.  17,  p.  211,  1905. 


THE  RELATION  OF  DIET  TO  DISEASE       425 

majority  among  the  rarer  specimens,  ten  carnivorous, 
seven  herbivorous  and  one  omnivorous  mammals,  ten 
carnivorous  and  two  seed-eating  birds,  due  possibly  to 
inappropriate  diet  or  to  some  unknown  factor  that  ren- 
dered the  diet  inadequate.  At  the  autopsy  nothing  was 
found  to  account  for  death  except  the  draining  of  all 
storage  supplies. 

Starvation". 

The  reports  of  studies  conducted  during  long  labora- 
tory fasts  have  been  among  the  most  valuable  records  for 
the  understanding  of  the  chemical  requirements  of  diet 
and  of  the  close  chemical  interrelationship  existing 
between  the  different  food  factors.  In  absolute  starva- 
tion life  is  very  short,  primarily  because  water  is 
necessary  for  respiration,  for  dissolving  products  of 
metabolism  and  for  preventing  changes  in  digestive  intes- 
tinal secretions.  The  amount  of  water  needed  varies  with 
different  species  of  animals.  If  the  water  is  supplied,  the 
organism  is  enabled  to  maintain  its  energy  for  continued 
existence  from  the  destruction  of  its  own  tissues.  The 
length  of  life  depends  upon  the  amount  of  protein 
ingested  before  the  fast  commenced,  and  the  amount  of 
stored  fat  and  glycogen,  especially  that  stored  in  the  liver./ 
The  mechanism  of  the  results  is  similar.  The  animal 
body  uses  first  its  available  glucose,  and  when  this  is 
partially  exhausted  burns  its  stored  fat  and  protein. 
The  fat  combustion  is  usually  defective,  ketone  bodies 
appearing  in  the  urine  in  large  quantities.  The  change 
from  fat  to  protein  metabolism  accounts  for  the  pre- 
mortal rise  in  metabolism  which  occurs  usually  a  few  days 
before  death.  The  chemical  composition  and  corpuscular 
richness  of  the  blood  is  tenaciously  preserved;  glucose 
and  protein  concentration  are  practically  normal  up  to 
the  day  of  death.  There  is  at  times  a  slight  increase  in 
globulins  and  always  an  increase  in  fat  due  to  its  trans- 
portation from  storage  depots.     The  cause  of  death  is 

28 


426  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

primarily  due  to  loss  of  substance  in  organs  necessary 
to  life  and  to  an  acid  intoxication. 

Wasting  occurs  first  in  stored  substances,  fat,  glyco- 
gen, etc.,  then  in  the  least  used  organs.  The  bones  usually 
show  some  rarefication.  The  animal,  as  a  rule,  dies  from 
acid  intoxication  before  atrophy  of  the  organs  is  marked.} 

In  the  wild,  when  animals  are  forced  to  seek  their  food 
with  the  expenditure  of  much  energy  and  where  feasts  are 
often  followed  by  fasts,  this  using  up  of  storage  supplies 

Table  20. 

Detailed  Analysis  of  the  Various  Diets  Used  at  the  Philadelphia  Garden  on 

Basis  of  100  Grams  of  Mixed  Food. 


Omnivora 

Carnivora 

Herbivora    Succu- 
lent Vegetables 

Herbivora  Coarse 
Food 

S 
B 

1 

1 

1 

S 

a 

0! 

1 

pq 

1 

>. 

K 

1 
1 

Protein 

Fat. 

14.3 
9.5 

26.7 
.034 
.058 
.497 
.103 
.263 
.117 
.338 
.0032 

11.5 
7.2 

41.2 
.068 
.093 
.713 
.284 
.484 
.377 
.486 
.0063 

15.6 

18.8 

.058 

.118 
1.694 

.421 
1.078 

.378 
1.146 

.015 

17.2 
.3 

.109 

.133 
1.671 

.373 
1.148 

.528 
1.119 

.0055 

6.1 

2.6 
18.5 
.067 
.164 
.538 
.08 
.556 
.038 
.134 
.0018 

3.2 
.5 
25.7 
.025 
.119 
.242 
.291 
.342 
.044 
.125 
.0012 

6.4 

2.2 
35.9 
.071 
.289 
.644 
.089 
.692 
.073 
.217 
.0022 

7.1 
1.3 

Carbohydrate 

Calcium 

Magnesium.  . 
Potassium .  .  . 

Sodium 

Phosphorus.  . 

Chlorin 

Sulphur 

Iron 

51.2 
.044 
.16 
.324 
.261 
.458 
.063 
.163 
.0012 

cannot  help  being  a  factor  in  preserving  the  integrity  of 
the  storage  and  eliminative  organs.  In  captivity  this 
cannot  occur.  Food  is  supplied  regularly,  exercise  is 
lacking,  consequently  overloading  and  disease  of  storage 
and  eliminative  organs  is  more  or  less  constant — a  situa- 
tion very  marked  in  the  Carnivora. 

A  further  study  of  Table  19  in  the  light  of  the  finer 
analysis  of  the  ingredients  of  the  diets,  shown  in  Table  20, 
explains,  at  least  in  part,  the  high  percentage  of  certain 
types  of  disease  in  relation  to  particular  diets. 

In  the  food  of  the  first  group,  the  omnivorous  mam- 
mals, there  is  a  moderately  increased  carbohydrate  con- 
tent and  an  unevenly  balanced  inorganic  content,  the 


THE  RELATION  OF  DIET  TO  DISEASE       427 

last  being  the  factor  most  at  fault.  The  calcium  and 
phosphorus  are  both  so  low  that  at  the  best  the  animal 
could  only  be  in  equilibrium,  while  any  drain  of  the  fixed 
bases  would  sooner  or  later  have  to  be  replenished  from 
the  calcium  and  phosphorus  storage  depots,  the  bones. 
Osteomalacia  is  most  marked  in  the  Cebidae,  monkeys 
whose  diet  is  even  lower  in  these  same  elements :  calcium 
.025,  phosphorus  .116,  and  iron  .0008  per  100  grams  of 
food.  The  inorangic  composition  of  all  animals  is  grossly 
similar ;  the  typical  digestion  developed  from  the  habitual 
diet  of  the  animal  explains  the  more  apparent  changes 
and  variations  in  their  reactions  to  certain  deprivations. 

Irkegularities  of  Inorganic  Metabolism. 

(Twelve  essential  elements  are  present  in  the  body, 
namely:  carbon,  nitrogen,  hydrogen,  oxygen,  phos- 
phorus, calcium,  sulphur,  sodium,  chlorine,  potas- 
sium, iron,  magnesium.  Of  these,  five  are  furnished 
by  the  protein  molecule  and  three  of  the  five  are 
duplicated  in  the  fats  and  carbohydrates;  the  remain- 
ing seven  must  be  present  in  the  mineral  ash.  These 
elements  functionate  in  three  ways,  (1)  as  constituents 
of  bone,  (2)  as  essential  elements  of  organic  com- 
pounds, (3)  as  soluble  salts  in  body  fluids.  Chlorine, 
sodium,  sulphur  are  supplied  in  sufficient  quantity  with 
most  diets.  In  the  case  of  chlorine,  marked  differences 
exist  between  the  herbivores  and  carnivores.  The  meat- 
eating  mammals  easily  acquire  sufficient  sodium  chloride 
from  the  flesh  and  blood  of  their  victims,  while  the 
herbivores  on  the  other  hand,  find  in  their  vegetable  food 
large  amounts  of  potassium  and  very  little  sodium  or^ 
chlorine  which  must  therefore  be  acquired  separately.) 
Both  omnivores  and  herbivores  crave  salt,  probably 
because  this  large  potassium  content  of  vegetable  food 
tends  to  increase  the  sodium  elimination.  A  deprivation 
of  salt  always  leads  to  a  distaste  for  foods  rich  in  po- 
tassium. So  far  as  is  known  excessive  sodium  stimulates 


428  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

protein  catabolisra,  and  through  the  overstimulation  of 
the  digestive  tract,  may  interfere  with  the  absorption 
of  food. 

Sulphur  is  largely  taken  into  the  body  in  organic 
combination  ^vith  the  protein,  (a  very  little  inorganic 
sulphur  appears  in  the  drinking  water)  therefore  if  the 
protein  requirements  are  adequate  the  sulphur  will 
usually  be  adequate. 

Magnesium  is  abundant  in  meat  and  most  plant 
tissues ;  so  that  except  in  diets  of  highly  refined  foods,  it 
is  more  often  excessive  than  deficient. 

The  other  elements,  calcium,  phosphorus  and  iron  are 
frequently  insufficient,  especially  for  animals  on  omniv- 
orous diet  (cf.  Table  20).  Phosphorus  enters  into 
every  living  cell,  and  in  cases  of  starvation  is  excreted 
up  to  the  last.  It  is  involved  in  practically  all  the  cell 
functions.  In  the  body  it  is  present  (1)  as  an  inorganic 
compound  in  the  bone  tissues  and  blood  where  it  helps  to 
maintain  neutrality,  (2)  as  phosphorus-containing  pro- 
tein, phosphatids  and  phosphoric  esters  of  a  carbohydrate, 
all  closely  associated  with  the  cell  and  its  nucleus.  In 
foods,  phosphorus  occurs  in  the  same  positions,  that  is, 
inorganically  or  combined  with  protein,  fat  or  carbohy- 
drate. It  is  not  entirely  proved  but  is  very  probable  that 
the  phosphorus  in  organic  combination  has  the  greater 
metabolic  value,  inasmuch  as  there  is  greater  storage  of 
nitrogen  and  stimulation  of  tissue  growth  on  foods  con- 
taining phosphorized  proteins,  fats,  etc.  It  has  been 
shown,  however,  that  the  animal  body  can  satisfactorily 
supply  its  phosphorus  requirements  by  inorganic  phos- 
phates. /The  omnivorous  diet,  even  the  widely  varied  diet 
of  man,  is  very  often  deficient  in  phosphorus,  a  fact  which 
becomes  very  important  when  we  consider  that  the 
omnivorous  diet  produces  many  acid  residues  wliich  must 
be  neutralized,  and  that  phosphorus  is  largely  responsible 
for  the  maintenance  of  tissue  neutrality.  Voit  showed 
that   the  phosphates   excreted   during  starvation  were 


THE  RELATION  OF  DIET  TO  DISEASE       429 

withdraAvii  from  the  bones ;  and  there  is  much  proof  that 
during  the  daily  metabolism  a  certain  slight  movement  of 
phosphorus  takes  place.  The  metabolized  phosphorus  is 
excreted  by  carnivores  practically  from  the  kidney  alone ; 
by  herbivores  almost  entirely  through  the  intestinal  wall, 
while  in  the  omnivores  it  is  excreted  by  kidney  and  intes- 
tinal tract.  Whether  these  facts  have  any  real  influence 
on  the  phosphorus  need  of  different  types  is  not  alto- 
gether determined. 

Calcium  also  enters  into  many  of  the  essential  func- 
tions of  life,  coagulation  of  the  blood,  contractility  of  the 
heart,  etc.  Omnivorous  diet  is  usually  deficient  in  this 
element,  which  is  very  irregularly  distributed  both  in 
animal  bodies  and  plants.  Insufficient  amounts  lead  to 
deprivation  of  body  tissues  and  to  the  production  of 
osteomalacia-like  conditions.  Voit  produced  marked  thin- 
ning of  the  skull  bones  and  sternum  by  a  diet  poor  in 
calcium.  Steenbok  and  his  associates  had  the  same 
results  in  cattle  by  feeding  ' '  shorts  ' '  a  diet  rich  in  mag- 
nesium. Etienne  (4)  showed  that  an  excess  of  magnesium 
in  an  otherwise  well  balanced  food  caused  a  continual 
loss  of  calcium.  Adults  stand  a  deprivation  of  calcium 
much  better  than  children  or  young  animals.  They  often 
show  no  symptoms  and  retain  a  normal  blood  content  as 
the  losses  from  the  blood  and  soft  tissues  are  promptly 
replaced  from  the  bones.  Sooner  or  later  all  these 
animals  show  weakness  and  flexibility  of  the  bones. 
Osteomalacia  occurred  in  5.2  per  cent,  of  the  animals  on 
an  omnivorous  diet,  that  is  this  number  showed  gross 
evidence  of  absorption  of  bone  salts.  This  condition 
occurring  in  man  and  the  lower  animals  is  a  generalized 
softening  of  adult  bones  that  were  at  one  time  normally 
calcified.  Three  clinical  varieties  are  recognized  in  man : 
a  mild  form  seen  in  pregnant,  puerperal  and  lactating 
women,  a  senile  form  in  which  the  lesions  are  usually 
limited  to  the  peh^s,   and  a  severe  progressive  form 

(4)   Jour.  Physiologic  et   Path.,  Vol.  14,  108,  1912. 


430  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

encountered  in  both  sexes  and  at  any  age.  This  last  form 
ends  in  marasmus.  Its  chemical  characteristic  is  a  loss 
of  calcium  and  phosphorus  with  retention  of  sulphur 
and  magnesium. 

The  progressive  type  has  occurred  very  frequently 
among  the  CebidaB  whose  diet  on  careful  examination, 
showed  a  protein  content  low  in  quantity,  poor  in  quality, 
and  especially  deficient  in  the  phosphorus-containing  pro- 
teins and  total  fat.  The  carbohydrate  was  high.  The  ash 
was  small  in  amount  and  predominatingly  acid.  The  daily 
ration  often  showed  only  an  unweighable  trace  of  calcium, 
phosphorus  or  iron.  Sodium,  potassium,  sulphur  and 
magnesium,  on  the  contrary,  were  present  in  amounts 
sufficient  for  equilibrium  or  in  excess.  The  Vitamines 
A.B.G.  were  present  but  were  not  always  correctly  pro- 
portioned. The  fat  soluble  A  was  low  and  in  some  daily 
rations  was  entirely  lacking. 

Diet  has  at  various  times  been  proposed  as  at  least 
one  factor  in  the  production  of  this  condition,  a  premise 
that  has  gained  considerable  weight  through  the  increase 
in  the  number  of  cases,  both  in  man  and  in  the  domesti- 
cated animals,  reported  from  the  war-famine  district  of 
Central  Europe  where  the  dietary  was  restricted  and 
unbalanced.  It  has  been  shown  that  w^hen  calcium  is  low 
in  the  diet,  the  amount  excreted  materially  exceeds  the 
intake.  Benedict  (5)  has  further  shown  that  even  during 
absolute  fasts  calcium  is  excreted.  The  requirements  of 
this  element  for  man  have  been  fairly  well  worked  out, 
but  for  animals  we  have  no  standards.  Still  it  seems 
certain  from  the  foregoing  observations  that  storage 
supplies  are  called  upon  very  early  in  cases  of  depriva- 
tion, while  in  pregnancy  and  lactation  when  the  calcium 
requirements  are  greatly  increased,  a  reason  is  found  for 
a  higher  incidence  of  osteomalacia.  Steenbok  and  Hart  (6) 
have  shown  that  the  skeletons  of  cows  and  goats  gave 

(.5)    Carnegie  Institute  Publication,  No.  203,  p.  247,  1915. 
(6)   Jour.  Biol.  Chem.,  Vol.  14,  p.  59,  1913. 


THE  RELATION  OF  DIET  TO  DISEASE       431 

evidence  of  a  drain  of  inorganic  salts  during  the  produc- 
tion of  milk  unless  the  calcium  and  phosphorus  of  the 
diet  were  liberally  supplied.  In  osteomalacia  it  would 
seem  that  inefficient  diet,  if  not  the  cause,  was  at  least  a 
very  potent  factor  in  pathogenesis.  The  disturbance 
of  the  calcium-phosphorus-metabolism  may  be  due  to 
the  deprivation  of  the  alkaline  salts  as  in  the  famine 
osteomalacia,  to  a  drain  from  the  alkaline  storage  of  the 
body  associated  with  an  inefficient  diet  as  in  the  osteo- 
malacia of  pregnancy  and  lactation  or  to  the  combined 
action  of  a  diet  faulty  in  more  than  its  salt  content,  which 
by  the  production  of  acid  in  its  oxidation  and  by  favoring 
the  development  of  acid-forming  bacteria,  causes  a  drain 
of  the  body  alkali  for  neutralization  of  the  acid ;  or  it  may 
possibly  be  due  to  a  combination  of  all  these  factors  act- 
ing through  their  influence  on  the  ductless  glands. 

Paget 's  disease  or  Osteitis  Deformans  is  a  chronic 
constitutional  process  which  usually  involves  all  the 
bones  of  the  adult  skeleton.  DaCosta  (7)  believed  it  to  be 
a  disorder  of  bone  metabolism  probably  dependent  upon 
absence  or  perversion  of  some  internal  secretion.  We 
have  had  the  unique  opportunity  of  observing  three  cases 
of  this  disease  in  Cebidse,  the  family  of  monkeys  which  has 
presented  the  highest  incidence  of  osteomalacia.  The 
experience  is  all  the  more  interesting  because  of  the 
typical  picture  presented  by  the  specimens,  and  of  the 
absence  of  references  in  the  literature  on  the  subject,  to 
the  occurrence  of  the  malady  in  wild  animals.  The  inter- 
esting point  about  these  cases  lies  in  the  fact  that  the 
disease  appeared  in  all  three  only  after  lime  water  was 
added  to  the  diet  to  supply  the  deficiency  of  calcium. 

Search  for  literary  record  of  the  disease  brought  to 
light  a  case  in  a  horse  that  Barthelemy  (8)  described,  but 
this  involved  the  epiphyses  of  the  bone  while  osteitis 

(7)  Publication  of  the  Jefferson  Medical  College  and  Hospital,  Vol. 
6,  p.  1,  1915. 

(8)  These  de  Lyon,  1901. 


432  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

deformans  is  confined  as  a  rule  to  the  diaphyses.  This 
case  was  probably  more  closely  allied  to  osteitis  fibrosa 
cystica.  Goldman  (9)  described  examples  in  fowls  and 
Rossweg  (10)  refers  to  specimens  in  domestic  goats  and 
monkeys.  Many  of  these  cases  first  come  under  obser- 
vation through  fractures,  an  accident  common  to  osteo- 
malacia, but  very  rare  in  well  developed  osteitis 
deformans.  The  diet  of  our  monkeys  was  exceeding  low 
in  those  substances  essential  to  bone  development. 
Sherman  (11)  has  shown  that  the  calcium  balance  is  reg- 
ulated to  a  certain  extent  by  the  calcium  ingested,  and 
that  when  the  diet  is  poor  in  this  element,  the  output 
materially  exceeds  the  intake,  a  condition  which  is 
definitely  changed  when  the  animal  is  put  on  a  diet  liigh 
in  calcium. 

So  far  as  we  could  find  there  are  no  recorded  studies 
of  the  mineral  metabolism  of  beginning  cases  of  Paget 's 
disease.  It  seems  possible  from  the  study  of  osteomalacia 
that  the  low  mineral  and  otherwise  faulty  diet,  added  to 
the  symptoms  produced  by  that  diet  might  so  disturb  the 
chemical  equilibrium,  directly  through  the  neurotrophic 
mechanism  or  through  the  perversion  of  the  ductless 
glands,  that  the  mere  addition  of  the  lime  water  might 
entirely  change  the  pathological  picture.  This  is  in 
accord  with  the  histology  where  the  initial  lesion  is 
resorption  of  bone  followed  by  irregular  proliferation. 
It  is  also  in  accord  with  the  probable  chemistry  of  cal- 
cification. These  animals  all  showed  a  lowered  carbon- 
dioxide-carrying-power  of  the  blood,  and  therefore 
lowered  ability  to  carry  calcium  in  solution.  It  is  possible 
that  Paget 's  disease  is  but  a  stage  in  a  deficiency  disease, 
a  faulty  reparative  response  through  a  disordered  neuro- 
trophic mechanism,  or  through  a  perversion  of  the  glands 
governing  calcium  metabolism.     Such  perversion  could 

(9)    Verein  Freiburger  Aerzte,  May  30,  1902. 

(10)  Vet.  Med.  Inaug.  Diss.  Giessen,  1913. 

(11)  Chemistry  of  Food  and  Nutrition,  Macmillan,  1918. 


THE  RELATION  OF  DIET  TO  DISEASE       433 

be  caused  by  an  improperly  balanced  diet,  or  by  the 
addition  of  an  excess  of  calcium  to  the  diet  of  an  animal 
whose  body  fluids  were  unable  by  reason  of  previous 
faulty  diet  or  other  disorder,  to  hold  it  in  solution.  In 
young  animals  the  calcium  demands  are  much  higher  than 
in  adults,  a  need  met  in  the  high  calcium  content  of  breast 
milk,  a  content  in  excess  of  almost  every  other  food,  but 
apparently  just  sufficient  to  maintain  calcium  equilibrium. 
After  it  is  weaned  the  young  animal  frequently  shows 
disorders  of  its  inorganic  metabolism.  Herter  estimated 
that  a  child  should  store  at  least  0.1  gram  of  calcium  daily 
and  he  described  many  cases  of  arrested  bone  develop- 
ment occurring  during  infancy  and  early  childhood, 
because  of  an  inefficient  assimilation  of  calcium.  One 
case,  probably  of  this  character,  was  found  in  a  Hama- 
dryas  Baboon  (Papio  hamadryas)  a  typical  example  of 
infantilism.  The  animal  was  an  adult  male  about  half 
the  size  of  an  adult  female.  His  skin  was  fine  and  more 
delicate  than  normal,  the  bones  were  small  and  slender, 
contour  of  body  was  that  of  a  young  animal,  genitalia 
were  imperfectly  developed,  thyroid  gland  apparently 
normal,  gastrointestinal  tract  atrophic,  associated  was 
a  slight  arthritis,  portal  cirrhosis  of  liver  and 
diffuse  nephritis. 

First  among  the  results  of  inorganic  insufficiency  in 
youth  stands  Rickets.  This  disease  occurs  in  children 
starting  usually  at  about  the  sixth  month  and  continuing 
with  irregular  remissions  for  several  years.  The  bone 
changes,  which  are  the  most  prominent,  are  always 
associated  with  more  or  less  severe  anemia,  a  general 
lowered  resistance  and  flabby  musculature.  The  excre- 
tion of  calcium  is  very  high  in  the  feces  and  low  in  the 
urine.  There  is  a  frequent  negative  calcium  balance 
dependent  upon  the  great  loss  in  the  feces.  Healing  is 
preceded  by  a  hyperretention  of  calicum  and  a  relative 
increase  in  the  urinary  calcium.  The  excessive  loss  of 
calcium  in  the  feces  is  not  brought  about  through  the 


434   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

agency  of  fats  because  fat  could  only  remove  calcium  as 
insoluble  soaps  and  these  are  not  at  all  increased.  This 
fact  contradicts  the  idea  of  fat  starvation  as  a  cause  of 
rickets.  Rowland  and  Kramer  found  that  the  blood  in 
active  rickets  bad  a  normal  or  slightly  lowered  calcium 
content,  but  a  regularly  reduced  phosphorus  content. 
The  latter  deficiency  was  extreme  at  times.  They  ascribe 
to  this  deficiency  the  failure  of  the  bones  to  calcify.  It 
can  be  readily  understood  that  a  decrease  of  phosphorus 
in  the  blood  would  render  difficult  the  precipitation  of 
calcium  phosphate. 

Recently  two  series  of  studies,  the  first  by  Pappen- 
heimer,  Zucher  and  McCann  and  the  second  by  Shipley, 
McCollum,  Park  and  Simonds  have  shown  that  rats  fed 
on  a  diet  low  in  calcium  but  with  a  sufficient  amount  of 
fat-soluble  vitamine  and  phosphorus  develop  a  bone  con- 
dition with  many  fundamental  resemblances  to  rickets. 
They  were  also  able  to  produce  the  condition  with  an 
excess  of  calcium  and  deficiency  of  phosphorus.  On  the 
first  diet,  the  condition  differs  from  rickets  in  that  the 
arrangement  of  the  proliferating  zone  of  cartilage  cells 
is  maintained  and  the  evidence  of  bone  resorption  in  the 
diaphyses  is  excessive.  A  diet  deficient  in  both  calcium 
and  phosphorus  leads  to  an  atypical  rickets. 

In  the  animals  autopsied  at  this  Garden  rickets 
occurred  very  much  more  frequently  in  the  flesh-eaters 
than  in  any  of  the  other  dietary  groups.  On  closer 
analysis  it  was  found  that  rickets  in  almost  every  case 
appeared  in  the  carnivores  which  did  not  receive  bones 
as  a  part  of  the  food.  Rickets  occurred  frequently  in  the 
omnivorous  macaques  which  however  did  not  show  osteo- 
malacia, although  they  belong  to  the  same  dietary  group 
as  the  Cebid^e.  The  reason  they  did  not  suffer  the  latter 
disease  while  adult  but  had  rachitic  young  is  probably 
due  to  the  fact  that  this  monkey  group,  which  breeds  best 
in  our  Garden,  receives  in  addition  to  the  diet  given  to 
CebidaB  one  raw  egg.    This  increased  the  calcium  content 


THE  RELATION  OF  DIET  TO  DISEASE       435 

of  their  food  more  nearly  to  the  requirements  of  these 
mammals.  These  monkeys  also  have  mouth  sacs,  which 
enable  them  to  acquire  more  food  per  Mlo  of  body  weight 
than  the  smaller  Cebidae  which  are  not  so  advantageously 
equipped.  The  food  even  in  the  amounts  consumed  by 
the  macaques  is  low  in  calcium,  phosphorus  and  iron.  It 
is  verj^  possible  that  there  are  enough  of  these  ingredients 
present  as  a  rule,  to  maintain  the  animal  in  organic 
equilibrium,  during  normal  life,  and  possibly  even  enough 
to  supply  the  needs  of  the  embryo  but  not  sufficient  to 
maintain  the  young  during  the  period  of  lactation.  A 
few  macaques  dying  during  the  delivery  of  young  showed 
slight  osteomalacic  changes  in  the  pelvis.  This  was 
notably  present  in  one  described  in  detail  by 
E.  A.  Schumann. 

The  calcium  requirements  of  the  female  are  always 
much  increased  during  pregnancy  and  lactation  due  to 
the  withdrawal  from  the  mother  to  meet  the  needs  of  the 
embryo  and  nursling.  Forbes  and  Beegle(12)  found 
that  lactating  animals  made  heavy  drains  on  their  stor- 
age calcium  even  when  the  diet  was  liberal  and  the  animal 
was  storing  nitrogen. 

Iron  is  the  essential  element  of  hemoglobin  and 
chromatin — the  body  constituent  most  directly  concerned 
with  the  process  of  oxidation,  secretion,  reproduction 
and  development.  The  iron  of  the  food  is  absorbed  from 
the  small  intestines,  enters  the  circulation  through  the 
l}Tnphatics,  is  deposited  in  the  liver,  spleen,  and  bone 
marrow  and  eliminated  through  the  intestinal  walls. 
There  is  very  little  iron  reserve  in  the  adult  body;  and  as 
a  result  any  failure  of  the  intake  to  equal  the  output 
causes  an  immediate  reduction  of  the  hemoglobin.  Voit 
found  that  the  iron  eliminated  in  the  feces  of  starving 
dogs,  or  dogs  on  a  diet  low  in  iron  comes  from  the  body 
through  the  intestinal  walls.  Medicinal  iron  stimulates 
the  production  of  hemoglobin  and  red  blood  cells  but 

(12)    Ohio  Agricultural  Experiment  Station  Bull.,  295. 


436   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

whether  it  is  directly  employed  in  the  production  of  hemo- 
globins has  not  been  proved.  Undoubtedly  most  of  the 
extra  iron  given  ^vith  the  food  passes  through  the  ali- 
mentaiy  tract  ^vithout  being  absorbed  or  metabolized. 
The  greater  the  amount  of  iron  in  the  food,  the  greater  the 
influence  of  the  inorganic  iron.  Anemia  occurred  in  all 
the  animals  we  examined  at  least  four  times  as  frequently 
in  omnivorous  as  in  all  the  other  dietary  groups,  a  fact 
probably  explained  by  the  low  content  of  iron  and  calcium 
in  this  diet.  Both  Von  Wendt(13)  and  Sherman  (14) 
demonstrated  that  larger  amounts  of  iron  were  required 
to  maintain  the  iron  equilibrium  when  the  amount  of  cal- 
cium was  low. 

Herter  has  sho^vn  that  many  anemias  are  associated 
with  intestinal  putrefaction.  The  carnivores,  however  on 
a  diet  that  putrefies  very  easily  and  on  one  in  which  the 
iron  content  is  apparently  of  distinctly  lower  nutritive 
value  than  that  of  the  iron  found  in  milk,  eggs  and  vege- 
tables, presented  an  anemic  incidence  of  only  0.32  per 
cent.  Tliis  is  probably  due  to  the  excellent  hygienic  care 
of  the  meat  foods  and  to  the  morphology  of  the  carniv- 
orous intestinal  tract,  which  is  short,  straight  and 
fashioned  for  quick  elimination.  The  cases  of  anemia 
steadily  increase  among  the  animals  as  the  conformation 
of  the  tract  approaches  the  omnivorous  type  with  the 
longer  and  wider  hind-gut. 

Herbivora,  obtaining  their  iron  from  vegetable 
sources,  are  much  less  liable  to  blood  disorders.  The 
iron  needs  of  the  female  are  greater  than  those  of  the 
male  because  of  the  drains  of  pregnancy  and  lactation. 
Young  animals  demand  more  iron  than  adults.  All 
exclusively  breast-feeding  animals  have  a  considerable 
storage  of  iron  in  the  body  at  birth,  while  those  that  eat 
food  immediately  have  no   such  supply.     Bunge's(15) 

(13)  Skandinavisches  Archiv.  f.  Physiologie,  Vol.  17,  p.  211,  1905. 

(14)  Bull,  185,  Experiment  Station,  U.  S.  Dept.  Agriculture,  1907. 

(15)  Physiological  and  Pathological  Chemistry,  Blakiston,  1902. 


THE  RELATION  OF  DIET  TO  DISEASE       437 

experiments  showed  that  breast-fed  animals  contained 
about  six  times  as  much  iron  as  the  milk  that  nourished 
them.  The  iron  content  of  all  these  animals  is  lughest 
at  birth,  remains  constant  during  the  suckling  period  and 
then  rapidly  decreases  to  the  adult  standard.  After  this 
level  is  reached  the  iron  metabolized  must  be  supplied 
from  the  food  if  the  hemoglobin  is  to  be  spared. 

The  functions  of  all  these  inorganic  substances  are 
intimately  interrelated  and  in  places  interchangeable. 
Calcium  is  capable  of  correcting  disturbances  of  inor- 
ganic equilibrium  in  the  animal  body  whatever  the  direc- 
tion of  the  deviation  from  the  normal  may  be.  These 
interrelationships  are  most  involved  in  the  maintenance 
of  body  neutrality.  The  normal  processes  of  metabolism 
involve  a  continual  production  of  carbonic,  phosphoric 
and  sulphuric  acid  which  must  be  immediately  disposed 
of  if  the  neutrality  of  the  body  is  to  be  permanent. 

The  factors  involved  in  this  are  carbonates,  phos- 
phates, ammonia  and  proteins.  Carbon  dioxide  is  the 
chief  excretory  product  but  is  at  the  same  time  a  normal 
constituent  of  the  blood  and  as  such,  is  an  important 
factor  in  this  physicochemical  reg-ulation.  There  is  a 
tendency  for  the  respiratory  mechanism  to  hold  its  carbon 
dioxide  tension  nearly  constant.  Late  investigations  have 
sho^m  that  lowering  of  this  tension  is  an  early  sign  of 
beginning  acidosis.  When  food  such  as  protein,  is  taken 
in  excess  the  strongly  acid  residues  are  neutralized  by 
the  sodium  and  potassium  carbonates  which  are  elimi- 
nated with  a  corresponding  loss  of  sodium  and  potassium. 
The  carbon  dioxide  tension  diminishes,  37.2  per  cent,  on  a 
high  protein  as  against  43.3  per  cent,  on  a  vegetable  diet. 
If  this  excess  is  long  continued,  the  result  may  be,  and 
often  is,  an  increased  elimination  of  the  base-forming  ele- 
ments which  if  not  made  good  tends  to  diminish  the  body's 
reserve  alkalinity.  A  diet  with  a  preponderance  of  basic 
elements  leads  to  an  alkaline  urine  with  an  increased  uric 
acid  solvency  and  an  increased  carbon  dioxide  tension 


438   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

and  reserve  alkalinity.  A  diet  with  a  preponderance  in 
the  acid-forming  elements,  on  the  contrary,  leads  to  an 
increased  urinary  acidity  and  urinary  ammonia, 
decreased  ability  to  dissolve  uric  acid  and  lowered  car- 
bon dioxide  tension  and  alkaline  reserve. 

Deficiencies  of  Vitamines, 

Recent  investigations  have  shown  that  diets  furnish- 
ing sufficient  amounts  of  protein,  fat,  carbohydrate  and 
inorganic  salts  may  yet  prove  inadequate  for  growth  or 
even  for  maintenance.  Hopkins,  (16)  feeding  rats  on  puri- 
fied food  mixture  was  unable  to  obtain  any  growth  until 
he  added  small  quantities  of  milk  or  of  the  ether-soluble 
portion  of  milk  but  with  this  addition  growth  progressed 
in  the  normal  manner,  but  it  was  out  of  all  proportion 
to  the  energy  or  protein  value  of  the  addition.  Five 
substances  of  this  character,  called  by  Funk  (17)  Vita- 
mines,  have  been  described,  two  of  wliich  have  definitely 
established  a  place  as  essential  food  factors.  According 
to  him,  pellagra,  rickets,  scurvy  and  beriberi  are  the 
result  of  a  lack  of  these  unidentified  but  specific 
and  indispensable  food  complexes. 

The  first  vitamine  isolated  was  the  fat  soluble  A,  an 
adequate  supply  of  which  is  necessary,  not  only  because 
of  its  stimulating  growth  properties,  but  because  its 
absence  produces  a  serious  condition  of  the  eyes  and,  at 
times,  marasmus  leading  to  death.  Xerophthalmia  is  a 
common  condition  in  animals  on  experimental  diets.  The 
eyes  are  swollen,  the  cornea  inflamed  and  often  opaque 
while  blindness  and  death  invariably  occur  unless  the  die- 
tary error  is  corrected.  McCollum(18)  rescued  animals 
almost  at  the  point  of  death  by  butter  or  other  fat  rich  in 
this  vitamine.  Opacities  of  the  cornea  are  often  seen  in  the 
animals  in  this  and  other  gardens  among  ungulates — hay- 

(16)  Joum.  Physiol.,  1912,  XLIV,  425. 

(17)  Die  Vitamine  und  ihre  Bedeutung  fur  die  Physiologie  und  Pa- 
thologic mit  besonderer  Beriicksichtigung  der  Avitaminoses,  Wiesbaden,  1914. 

(18)  Newer  Knowledge  of  Nutrition,  Macmillan,  1919. 


THE  RELATION  OF  DIET  TO  DISEASE       439 

eating  mammals ;  four  advanced  cases  were  found,  three 
in  seed-eating  birds  and  one  in  a  fox  on  a  diet  made  up 
solely  of  horse  muscle.  The  quantity  of  vitamine  A 
present  in  muscle,  hay  and  seeds  is  very  small.  It  is 
supplied  in  largest  amounts  in  milk,  eggs,  glandular 
organs  and  leaves,  substances  which  were  very  low  or 
absent  in  the  diet  of  all  the  affected  animals.  This 
xerophthalmia  has  been  reported  in  man  on  several  occa- 
sions, especially  by  Hrdlicka(19)  in  American  Indians, 
by  Mori (20)  in  1400  Japanese  during  a  period  of  food 
shortage  (this  epidemic  was  cured  by  the  addition  of 
chicken  livers  to  the  diet),  by  Bloch(21)  in  forty-seven 
children  of  Copenhagen  fed  on  a  fat  free  milk  who  were 
cured  by  the  administration  of  cod  liver  oil.  The  disease 
is  not  however  a  fat  starvation,  as  it  is  entirely  unin- 
fluenced by  vegetable  fats  which  do  not  contain 
this  Adtamine. 

Beriberi  is  an  established  deficiency  disease,  fre- 
quently seen  among  the  poorer  classes  of  the  Orient 
whose  diet  is  limited  to  polished  rice  and  fish.  It  has 
appeared  in  Labrador  coincident  with  the  excessive  use 
of  bolted  flour.  A  similar  condition  has  been  induced  in 
pigs  and  cattle  by  a  diet  made  up  of  an  excess  of  cotton 
seed  meal  and  tankage.  Two  forms  of  the  disease  are 
described:  (1)  acute  or  wet,  characterized  by  marked 
edema,  ascites,  hydropericardium,  hydrothorax,  edema  of 
the  lungs,  and  a  congestion  of  the  spleen,  liver,  kidney, 
and  heart  muscle,  (2)  chronic  or  dry,  characterized  by 
polyneuritis.  The  disease  was  first  produced  experimen- 
tally in  pigeons  by  Eijkman(22)  in  1897  by  means  of  a 
diet  of  polished  rice.  The  paralysis  appeared  in  2-3 
weeks  after  the  diet  was  initiated.  Eraser  and 
Stanton  (23)  in  1907,  found  that  it  could  be  cured  by  an 

(19)  B7ill.,  34,  Bureau  of  Amer.  Ethnolo^. 

(20)  Jahrbuch.  Kinderheilk.,  1904,  LIX,  175. 

(21)  Journ.  Am.  Med.  Assoc,  1917,  LXVIII,  1516. 

(22)  Arch.  Path.  Anat.,  1897,  CXLVIII,  523. 

(23)  Lancet,  London,  March  12,  1910,  733. 


440   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

alcoholic  extract  of  rice  polishings.  Funk (24)  later 
determined  the  vitamine  character  of  this  extract.  In 
pigeons  and  fowls  experimental  feeding  usually  results 
in  the  chronic  or  polyneuritic  form,  expressed  by  a  typical 
degenerative  inflammatory  condition  of  the  peripheral 
nerves.  In  pigs,  on  the  contrary,  Rommel  and 
Vedder(25)  produced  both  types,  though  the  acute  or 
wet  beriberi  appeared  more  frequently.  In  rats  the  same 
deficiency  causes  multiple  hemorrhages  in  the  cerebellum 
and  midbrain  followed  by  a  degeneration  of  the  associ- 
ated nervous  structures.  It  is  possible  that  the  pathol- 
ogy following  a  lack  of  the  vitamine  B  or  in  fact  any  of 
the  vitamines  will  vary  with  the  different  species  or 
with  varying  demands  of  different  individuals.  This 
antineuritic  vitamine  affects  more  than  the  nervous 
system,  and  it  is  possible  that  all  vitamines  may  have 
mder  effects  than  are  at  present  described. 

Scurvy  was  the  first  condition  to  call  attention  to  diet 
as  a  cause  of  disease.  It  occurs  in  man  when  deprived 
of  fresh  vegetables.  That  faulty  diet  was  in  some  way 
the  cause  of  scurvy  has  been  known  for  many  years, 
but  only  since  1905  has  there  been  any  systematic  attempt 
to  determine  the  peculiar  value  of  the  curative  foods.  At 
this  time  Theobald  Smith (26)  called  attention  to  a 
disease  suggestive  of  scurvy^  which  developed  in  guinea- 
pigs  fed  on  a  diet  of  oatmeal.  This  observation  was  con- 
firmed by  Hoist  and  Frolich(27)  who  stated  that  the 
disease  could  be  prevented  by  the  addition  of  fresh  milk 
or  cabbage,  because  in  these  foods  there  was  present  an 
antiscorbutic  or  C  vitamine.  This  unidentified  substance 
was  easily  destroyed  or  diminished  by  heat  or  an  alkaline 
medium.  It  was  found  in  rather  large  amounts  in 
succulent  vegetables  and  fruits.     McCollum(28)  and  his 

(24)  Lancet,  London,  1911,  II,  1266. 

(25)  Bull,  Dept.  of  Agriculture,  Dec.  13,  1915. 

(26)  Bureau  of  Animal  Industry,  1895-96,  172. 

(27)  Z.  Eyg.  u.  Infektionskrankh.,  1913,  LXXV,  334. 

(28)  Jour.  Biol.  Chem.,  1917,  XXXI,  229. 


THE  RELATION  OF  DIET  TO  DISEASE       441 

coworkers  showed  that  the  oat  kernel  was  low  in  inor- 
ganic salts  and  vitamine  A  and  poor  in  the  quality  of  its 
protein;  but  with  these  faults  corrected  it  proved  to  be 
a  complete  food  for  rats.  McCollum  also  found  that 
scurvy  developed  more  readily  in  animals  if  the  physical 
properties  of  the  diet  favored  constipation.  He  was  able 
to  delay  the  onset  of  the  disease  in  guinea-pigs  for  a 
considerable  period  by  the  addition  of  mineral  oil  which 
has  no  food  value,  or  phenolphthalein,  a  cathartic.  At 
the  same  time,  Jackson  and  Moore, (29)  found  the  cecum 
of  all  guinea-pigs  dying  of  scurvy,  packed  with  putrefying 
feces.  They  were  able  to  produce  a  mild  type  of  the  dis- 
ease by  the  injection  of  the  diplococci  isolated  from  the 
swollen  joints. 

From  these  observations  it  seems  safe  to  conclude  that 
scurvy  may  not  be  purely  a  deficiency  disease,  or  even  a 
simple  dietary  one,  although  the  presence  of  a  vitamine 
influence  is  not  excluded ;  but  it  is  probably  the  result  of 
a  bacterial  invasion  of  tissues  debilitated  by  a  faulty  diet 
and  by  the  toxins  produced  by  the  putrefactive  bacteria 
developing  in  a  diet  unsuited  to  the  anatomical  demands 
of  the  alimentary  tract.  This  theory  receives  support 
from  the  fact  that  pasteurization  destroys  all  aciduric 
bacteria,  allowing  only  the  spore-forming  putrefactors 
to  develop ;  and  from  the  fact  that  scurvy  develops  more 
frequently  in  children  on  stale  pasteurized  than  on  stale 
raw  or  boiled  milk.  In  this  Garden  no  suggestion  of  scurvy 
has  been  noted. 

Pellagra  is  very  definitely  a  disease  of  poverty  en- 
demic for  years  among  the  poor,  especially  in  the  moun- 
tains of  Northern  Italy,  It  has  been  under  observation 
in  the  United  States  since  1907.  So  far  as  is  known  no 
cases  have  been  observed  among  animals.  Opinions  dif- 
fer as  to  the  role  of  diet  in  the  etiolog}^  but  the  results 
of  recent  studies  seem  to  show  that  uncomplicated  cases 
of  average  severity  clear  up  entirely  on  a  diet  rich  in 

"         (29)   Jour.  Infect.  Dis.,  1916,  XIX,  478. 
29 


442   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

animal  protein.  No  vitamine  deficiency  has  so  far  been 
determined.  Wilson's  careful  studies  of  the  diets  known 
to  have  produced  the  condition  show  that  the  etiological 
factor  lies  in  a  deficiency  of  the  protein  molecule.  The 
results  of  Goldberger(30)  corroborate  this  fact,  and  he 
concludes  from  his  latest  studies  that  ''  the  dominating 
role  of  diet  in  the  prevention  and  causation  of  pellagra 
is  referable  primarily  to  the  character  of  the  protein  sup- 
ply or  to  the  specific  quality  of  the  aminoacid  makeup 
of  the  protein  supply. ' '  Just  what  aminoacid  or  combina- 
tion of  aminoacids  it  is,  has  not  been  determined,  nor  has 
the  possibility  of  a  vitamine  alone  or  in  combination  Avith 
the  aminoacid  factor  been  absolutely  excluded. 

The  principal  influence  of  the  omnivorous  diet  is 
toward  those  degenerations  arising  primarily  from  im- 
balances in  the  inorganic  makeup,  or  to  insufficiencies  of 
certain  necessary  factors.  The  vitamine  deficiencies  are 
markedly  less  prevalent  in  animals  than  in  man  whose 
food  is  less  often  consumed  in  its  natural  state.  It  is 
now  known  that  much  of  the  injury  and  loss  of  nutritive 
value  in  foods  is  produced  by  the  processes  involved  in 
preparation,  preservation,  refinement  and  storage. 
Whenever  the  choice  of  food  is  not  restricted,  vitamine 
deficiencies  do  not  occur.  The  vitamine  requirements 
probably  differ  in  different  species  and  in  individuals 
from  the  same  species  according  to  their  environmental 
and  individual  variations.  It  is  very  possible  that  if  the 
diet  is  low  in  vitamine  content  there  may  arise  conditions 
of  relative  deficiencies ;  and  McCarrison  has  sho^vn  that 
a  vitamine  deficiency  associated  with  a  high  fat  or  car- 
bohydrate content  may  disturb  the  balance  of  the  endo- 
crine glands.  It  is  however  to  the  inorganic  content  of 
the  omnivorous  food  that  most  of  the  disturbances  pecul- 
iar to  this  diet  are  to  be  assigned. 

With  the  flesh  eating  animals  and  birds  the  records 
present  a  very  different  picture.    Disorders  of  the  diges- 

(30)   Jour.  A.M.A.,  1922,  79,  2132.  " 


THE  RELATION  OF  DIET  TO  DISEASE       443 

tive  tube,  of  the  storage  organs,  of  the  organs  of  elimina- 
tion and  of  the  endocrine  glands  predominate.  Their 
diet  is  low  in  carbohydrates  and,  at  times,  in  fats  and  very- 
high  in  protein.  Bone  supplies  the  inorganic  salts,  which 
in  this  Garden  is  fed  only  to  the  larger  mammals.  The 
carnivorous  birds  get  their  inorganic  supply  from  mice 
which  are  eaten  entire.  The  carnivores  are  as  a  rule  large 
and  are  given  to  active  fighting  or  to  long  flights.  In  the 
mid,  very  probably  there  are  long  periods  between  feasts, 
while  in  captivity  the  food  is  always  plentiful  and  regu- 
larly supplied.  This  regularity  added  to  the  lack  of  exer- 
cise, particularly  among  the  larger  animals,  must  lead  to 
excessive  demands  upon  the  storage  and  eliminating 
organs.  Storage  is  always  promoted  by  rest  and  liberal 
diet,  and  cleared  away  by  exercise  and  starvation.  The 
life  of  these  birds  and  mammals,  moreover  favors  inac- 
tivity of  the  bowels,  which,  together  with  the  highly  putre- 
factive diet  adds  another  serious  factor  to  a  problem 
which  in  gardens  is  almost  insurmountable. 

Irregularities  of  Carbohydrate  Metabolism. 

The  carbohydrates  are  derived  from  the  glucose  and 
glycogen  of  the  meat  and  from  the  protein  molecule.  They 
are  absolutely  less  than  in  the  diet  of  herbivores  but  be- 
come a  factor  in  the  disorders  of  this  group  because  of 
the  lack  of  exercise  and  the  regularity  of  feeding.  In 
digestion  the  carbohydrate  becomes  available  for  absorp- 
tion and  bacterial  growth  in  the  upper  small  intestine 
and  appears  on  the  other  side  of  the  intestinal  wall  as 
blood  glucose  in  which  form  it  is  burned  for  energy  or 
stored  as  glycogen  for  the  future  maintenance  of  the 
blood  glucose. 

The  blood  of  different  animals  has  a  glucose  concentra- 
tion between  0.05  -0.1  which  for  each  species  is  quite  con- 
stant, as  it  is  regulated  by  the  coadaptation  of  four 
factors:  combustion,  fermentation  of  glycogen,  forma- 
tion of  fat,  and  elimination  from  the  kidney.    In  excessive 


444   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

feeding  the  amount  needed  for  energy  is  burned,  the 
remainder  is  stored  in  the  liver  up  to  its  capacity,  then 
in  the  muscles  and  other  cells,  after  which  fat  is  formed 
and  all  further  excess  is  eliminated  by  the  kidney.  Over- 
feeding causes  an  immediate  overloading  of  the  oxidative 
mechanism  mth  symptoms  of  gastric  disorder,  achylia, 
and  at  times  acid  fermentation  with  irritation  of  the 
stomach  walls  and  the  development  of  bacteria  in  the 
organ.  This  is  frequently  followed  by  glycosuria,  several 
types  of  which  are  described:  (1)  associated  \rith  an 
increased  concentration  of  glucose  following  excessive 
ingestion  exceeding  the  normal  glycogenic  function  of  the 
liver,  a  form  common  among  the  Herbivora,  (2)  that  due 
to  a  reduction  of  the  glycogenic  function  of  the  liver,  (3) 
that  associated  with  disease  of  the  ductless  glands  in 
which  the  resulting  glycosuria  probably  depends  upon  the 
influence  of  these  glands  upon  the  pancreas,  (4)  that 
dependent  upon  the  defect  of  glycolysis  or  to  an  over- 
stocked liver  seen  in  gout,  obesity  or  hypertrophic  cirrho- 
sis, and  (5)  renal  glycosuria  due  to  a  lowering  of  the 
renal  threshold  and  usually  associated  with  gout,  arterio- 
sclerosis or  chronic  nephritis ;  this  last  is  best  explained 
on  the  ground  of  increased  renal  permeability.  Normally 
when  the  blood  sugar  concentration  rises  above  a  certain 
level  the  elimination  via  the  kidney  begins  and  continues 
until  the  blood  has  again  reached  its  normal  concentration. 
The  relation  of  the  kidney  to  glucose  concentration  is 
not  constant  and  variation  is  always  toward  the  side  of 
lesser  elimination  while  the  kidneys  become  accustomed 
to  the  higher  level. 

Diabetes,  a  disease  of  the  islands  of  Langerhans  in 
the  pancreas,  is  essentially  a  disturbance  of  sugar  metab- 
olism always  associated  with  an  exaggerated  and  defec- 
tive fat  and  protein  combustion.  It  is  not  only  that  the 
diabetic  has  lost  the  faculty  of  combustion  but  these 
abnormalities  all  establish  states  of  intoxication  to  which 
the  diabetic  must  sooner  or  later  succumb.    Among  lower 


THE  RELATION  OF  DIET  TO  DISEASE       445 

animals  the  disease  is  rare.  Dogs  are  most  frequently 
affected  (about  1  in  12,000  deaths).  It  has  also  been 
described  in  horses,  cattle  and  monkeys.  In  our  records 
there  was  one  case  an  Artie  fox  {Canis  lagopus)  present- 
ing a  typical  picture.  Degeneration  of  the  islands  of 
Langerhans  was  seen  in  three  other  animals,  but  there 
was  no  other  evidence  of  diabetes.  This  disease  is  not 
due  to  diet  but  to  the  absence  of  a  normal  ferment  (pan- 
creaticozymo-excitor)  for  one  particular  type  of  food. 

iRREGULAErriES  OF   FaT  MeT ABOLISH. 

Disorders  of  fat  metabolism  are  very  rare  among 
lower  animals  notwithstanding  the  fact  that  fat  even  in 
the  carnivorous  diet,  represents  about  13  per  cent,  of  the 
whole  intake.  It  plays  two  important  roles  in  the  body, 
storage  for  energy  reserve,  and  as  a  most  essential 
structure  in  cellular  protoplasm,  in  which  position  it  joins 
with  protein  in  complex  combinations  of  still  unknown 
composition  which  present  to  a  striking  degree  the 
phenomenon  of  absorption.  Very  marked  biological 
differences  exist  in  the  value  of  fats  from  different 
sources,  due  to  the  presence  or  absence  of  vitamines. 
The  body  fat  is  derived  from  the  fat  of  the  diet  or  is 
synthesized  from  glucose.  The  former  is  specific  to  the 
fat  consumed  while  the  latter  is  specific  to  the  animal.  In 
omnivores  the  type  depends  upon  the  varying  extent  to 
which  animal  fats  enter  the  diet,  in  carnivores  it  depends 
almost  entirely  on  the  fat  intake,  while  in  the  herbivores 
practically  all  the  fat  is  synthesized  from  the  carbo- 
hydrate. On  digestion,  fat  splits,  yielding  a  glycerol  and 
fatty  acid  which  are  collected  in  the  lymph  spaces  of  the 
intestinal  mucosa,  there  changing  to  some  complex  com- 
bination which  is  not  only  soluble  but  diffusible. 

Fatty  infiltration  and  fatty  degeneration  are  condi- 
tions of  much  pathological  interest  and  of  great  frequency 
in  captive  animals.  The  researches  of  Mansfield (31)  have 

(31)   Pfluger's  Arch.,\^m  (129),  63. 


446   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

throAvn  considerable  light  upon  these  conditions.  He 
found  that  the  total  fat  content  in  cases  of  most  marked 
degeneration  was  normal  or  reduced.  The  proportion  of 
fat  free  from  protein  was  increased  and  the  firmly  bound 
fat  decreased.  This  increase  is  due  to  neutral  fat  brought 
from  without  the  organ  by  the  blood  when  for  any  cause 
the  oxidative  powers  are  decreased,  and  setting  free  of 
the  previously  invisible  intercellular  fat  and  lipoids, 
which  are  normally  present  in  the  cells,  by  autolytic  or 
physicochemical  changes.  This  condition  is  pretty  evenly 
distributed  among  the  dietary  groups,  the  liver  being 
most  commonly  involved.  The  hepatic  cells  are  easily 
degenerated  by  the  toxins  or  other  harmful  substances 
passing  through  the  organ  and  become  passive  and  unable 
to  throw  off  or  to  utilize  the  deposited  fat.  In  all  prob- 
ability the  same  general  situation  occurs  in  the 
atheromatous  changes  in  arteriosclerosis  which  on  this 
diet  shows  a  high  incidence.  The  causative  agent  is  prob- 
ably some  poisonous  substance,  possibly  a  protein 
degradation  product,  indol,  pressor  substance,  acting  on 
the  intima  over  long  periods,  or  at  irregular  but  often 
repeated  periods  causing  first  destruction  then  fat  accu- 
mulation. It  is  also  possible  that  it  may  be  caused  by 
repeated  absorption  of  some  sensitizing  protein. 
Arteriosclerosis  in  these  animals  is  often  closely  asso- 
ciated with  nephritis. 

Obesity  may  result  from  excessive  ingestion  of  food 
in  individuals  whose  habits  are  sedentary  and  whose 
digestions  are  active  or  it  may  come  from  an  inherent 
abnormality  of  metabolism  dependent  upon  ductless 
gland  disease.  It  is  very  common  in  castrated  animals. 
The  obesity  of  overeating  is  always  of  milder  type  than 
that  associated  with  endocrine  disturbance. 

So  far  as  is  known  there  are  two  main  disorders  of  fat 
metabolism — the  failure  of  the  diabetic  to  form  fat  from 
glucose,  and  acidosis,  the  inability  of  the  organism  to  burn 
fat  beyond  betaoxybutyric  acid,  acetoacetic  acid,  or  ace- 


THE  RELATION  OF  DIET  TO  DISEASE       447 

tone.  The  symptoms  are  unsteadiness  of  gait,  stupor, 
coma,  air  hunger,  in  all  of  which  the  essential  features  are 
due  to  the  impoverishment  of  the  body  in  available  bases. 
In  infants  this  frequently  follows  an  excessive  fat  diet. 
It  is  also  common  in  starvation  due  to  the  deprivation  of 
sugar.  It  is  associated  with  phosphorus  poisoning,  nar- 
cosis, carcinoma,  liver  disease,  inanition,  etc.  It  has  been 
produced  experimentally  by  the  administration  of  acids 
or  by  foods  deprived  of  their  bases.  The  excess  of  acid 
in  the  body  whether  produced  in  the  body  or  introduced 
from  without  must  be  neutralized  in  part  by  the  ammonia 
manufactured  in  the  ultimate  metabolic  transformation 
of  the  protein  and  by  the  alkaline  salts  of  the  blood  and 
tissues.  When  alkali  is  reduced  the  carbon  dioxide  accu- 
mulates in  the  tissues,  blocking  oxidation.  The  urine 
immediately  shows  an  increase  of  ammonium  salts,  a 
decrease  of  the  urea  and  an  increase  in  the  output  of 
sodium,  potassium,  calcium  and  magnesium,  which  last 
two  are  dra^vn  from  the  bones. 

Symptoms  do  not  arise  until  the  fixed  alkalies  are 
exhausted;  and  they  are  immediately  relieved  by  the 
administration  of  alkalies,  except  in  those  cases  of  starva- 
tion where  the  administration  of  sugar  and  the  subsequent 
sparing  of  the  fats  relieves  the  situation.  In  herbivores, 
acidosis  does  not  follow  starvation,  but,  on  the  other  hand, 
it  is  markedly  easier  to  excite  it  in  herbivores  than  in 
carnivores  whose  heavy  protein  diet  produces  more 
ammonia,  which  better  enables  the  animals  to  protect 
their  fixed  alkalies.  The  acid  intoxication  of  infections 
arises  from  different  causes  and  is  dependent  on  the 
intensity  of  the  type  of  infection;  but  ultimately  it  also 
depends  upon  the  depletion  of  the  fixed  alkalies. 

Ireegulaeities  of  Protein  Metabolism. 

Fat  and  carbohydrate  disturbances  are  not  infrequent 
in  carnivores,  but  it  is  with  the  protein  fraction  of  the  diet 
that  most  of  the  trouble  is  connected.     Natural  foods 


448   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

contain  several  proteins  or  groups  of  proteins,  whose 
biological  adequacy  depends  upon  their  yield  of  amino- 
acid.  Experiment  has  shown  that  many  proteins  are 
entirely  lacking  in  one  or  more  of  these  essential  radicles ; 
and  no  food  can  be  adequate  unless  it  contains  at  least 
all  the  aminoacids  that  the  individual  animal  is  unable  to 
manufacture  for  itself.  So  far  as  is  known,  no  animal  can 
produce  in  itself  either  lysin  or  tiyptophane.  Gliadin,  the 
principal  protein  of  wheat  and  lacking  in  lysin,  is  unable 
to  support  gro^\i:h  even  when  given  in  amounts  sufficient 
to  insure  the  storage  of  nitrogen,  and  is  associated  with  a 
diet  adequate  in  all  other  factors.  Absence  of  tryptophane 
prevents  not  only  growth  but  maintenance.  Any  of  the 
aminoacids,  whose  radicles  are  contained  in  tissue  pro- 
teins, may  contribute  to  the  maintenance  of  adult 
equilibrium ;  but  no  growth  occurs  unless  all  the  necessary 
groups  are  present.  Except  in  laboratories,  diets  are 
never  made  up  of  isolated  proteins,  but  they  are  often 
composed  of  proteins  derived  from  one  plant  and  are 
often  deficient.  McCollum  and  his  associates  in  their 
studies  showed  that  while  there  were  pronounced  differ- 
ences in  the  composition  of  many  foods  used  by  men  and 
animals  not  only  in  their  protein  content  but  in  water, 
fats,  carbohydrates,  etc.,  yet  in  the  combinations  found 
even  in  rather  restricted  diets,  the  errors,  as  a  rule,  cor- 
rected each  other. 

During  digestion  the  protein  molecule  is  broken  down 
into  the  component  aminoacids  which  are  absorbed  and 
synthesized  in  the  intestinal  walls,  and  appear  on  the 
other  side  as  the  specific  blood  protein,  which  serves  as 
the  substrate  for  the  anabolism  of  all  the  special  tissue 
proteins.  Excessive  protein  is  stored  to  a  slight  extent  as 
aminoacid  for  the  future  maintenance  of  the  blood  pro- 
tein, the  integrity  of  which  is  tenaciously  protected  during 
hibernations,  sexual  migrations,  and  even  during  starva- 
tion.    The   animal  body  tends   to   adjust  its   nitrogen 


THE  RELATION  OF  DIET  TO  DISEASE       449 

metabolism  to  its  nitrogen  supply;  the  adjustment 
requires  an  appreciable  amount  of  time.  A  diet  changed 
to  a  lower  nitrogen  level  results  in  a  continued  loss  of 
nitrogen,  increased  combustion  of  fats  and  carbohydrates. 
The  animal  makes  no  apparent  effort  to  reestablish 
equilibrium,  and  sooner  or  later  digestive  disturbances 
and  loss  of  strength  occur. 

If,  on  the  contrary,  the  protein  is  steadily  increased 
after  an  animal  has  established  equilibrium,  the  nitrogen 
metabolism  increases  and  the  level  of  nitrogen  equilibrium 
rises  to  higher  and  higher  levels.  There  is,  at  the  same  time, 
a  lowering  of  the  fat  combustion,  an  increase  in  the  respir- 
atory quotient  and  in  the  heat  production.  The  excess 
protein  must  be  split,  deaminated,  burned  and  eliminated. 
Fifty-five  per  cent,  of  the  intake  is  converted  into  glucose 
which  is  burned  and  the  excess  stored  as  glycogen.  The 
sulphuric  acid  formed  during  the  protein  cleavage  is 
neutralized  by  the  body  alkalies.  In  these  cases  the  liver 
is  often  congested  and  enlarged.  The  urine  shows  excess 
of  urea  and  ammonia.  At  times  the  excess,  being  so  great 
that  it  cannot  be  absorbed,  undergoes  chemical  and  bac- 
terial decomposition  which  causes  digestive  disturbances, 
torpor  and  constipation. 

The  organisms  associated  with  protein  food  are 
usually  the  putrefactive  types  which  break  the  protein 
molecule  into  the  aromatic  bodies,  phenols,  indolacetic 
acid,  indolpropionic  acid,  skatol,  etc.  These  bodies  on 
absorption  are  believed  to  give  rise  to  hypertrophy  of  the 
adrenal,  interstitial  changes  in  the  kidney,  and  arterio- 
sclerosis. Another  group  of  substances,  pressor  bases 
and  amines,  are  manufactured  by  certain  anaerobes 
acting  on  proteins.  These,  when  fed  by  mouth,  are  detoxi- 
cated  by  the  liver  cells,  but  when  formed  below  the  portal 
circulation,  give  rise  to  anaphylactic  phenomena — 
urticaria,  etc.  Certain  other  organisms  give  rise  to 
soluble  toxins  as  in  botulism  and  thyrotoxicon  poisoning. 


450  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

All  these  types  of  toxins  will  destroy  if  they  act  acutely  in 
sufficient  concentration;  or  as  is  more  common,  if  they  act 
persistently  over  long  periods,  or  at  oft  recurring  inter- 
vals they  will  cause  serious  injury  to  the  tissues  coming 
in  contact  with  them,  and  have  a  part  in  the  production 
of  cirrhosis  of  the  Hver,  chronic  nephritis,  myocarditis, 
arteriosclerosis,  etc. 

All  foods  have  a  limit  beyond  which  they  are  excreted 
untouched  or  imperfectly  oxidized.  Many  of  these  partial 
oxidation  products  of  protein  are  in  themselves  toxic  and 
may  also  be  a  source  of  these  degenerative  organ  condi- 
tions. The  pathological  material  studied  by  us  showed  a 
marked  decrease  in  gastrointestinal  diseases  in  close  asso- 
ciation with  the  more  hygienic  care  of  the  meat  foods. 

Always  associated  with  the  protein  foods  are  the 
nucleoprotein  complexes,  which  are  split  by  both  bacteria 
and  digestive  juices  into  globulins  and  nucleic  acid,  and 
then  through  the  agency  of  a  special  enzyme,  into  purin 
bases  and  uric  acid,  in  which  forms  they  are  excreted  in 
the  urine  and  feces.  The  oxidation  of  purins  is 
never  complete. 

Gout,  representing  the  pathology  of  purin  metabolism, 
is  a  paroxysmal  inflammatory  disturbance,  due  to  the 
deposition  of  sodium  urates  in  the  joints  or  in  the  internal 
organs,  usually  accompanied  by  a  fibrosis  especially  in 
the  liver,  kidney,  arteries,  etc.  The  disease  occurs  almost 
exclusively  in  birds.  Isolated  cases  have  been  described 
in  dogs,  horses  and  hogs,  but  among  lower  animals  it  is 
undoubtedly  very  rare.  In  birds  it  is  most  frequent  in  the 
carnivores — 4  per  cent.,  as  against  0.02  per  cent,  in  all 
other  groups.  It  is  higher  in  fish-eating  birds  than  among 
the  flesh-eaters.  The  avian  gout  is  usually  of  the  visceral 
type  and  was  most  marked  in  its  distribution  over  the 
organs  in  the  Anseres  and  Psittaci,  birds  whose  diet 
apparently  is  not  unduly  heavy  in  nucleoproteins,  but 
whose  tract  approaches  the  carnivorous  type.    The  only 


THE  RELATION  OF  DIET  TO  DISEASE       451 

arthritic  cases  occurred  in  Boatbilled  herons  {Cancroma 
cochlearia),  fish-eaters.  Our  records  show  examples  in 
Accipitres,  Galli  and  Columbae,  although  the  number  of 
cases  in  the  last  order  were  few  and  slight  in  extent.  Tliis 
disease  stands  in  close  relation  to  diet,  as  it  develops  on 
generous  protein  food,  high  in  nucleoprotein  or  hypo- 
xanthin,  especially  if  this  be  associated  with  restricted 
activity. 

The  carnivorous  mammals  lead  in  the  disease  of  the 
thyroid  glands.  Thyroid  disease  occurs  among  the  birds, 
but  is  equally  distributed  among  the  dietary  groups. 
Thyroid  activity  has  a  marked  influence  on  metabolism 
probably  through  the  influence  of  the  iodine-containing 
protein  of  its  secretion.  There  are  some  experimental 
evidences  in  favor  of  a  detoxicating  function  of  the  thy- 
roid, of  which  the  following  are  quoted :  (1)  The  effects  of 
thyroidectomy  are  most  marked  in  the  carnivores; 
Herbivora  are  often  capable  of  several  years  of  life  with- 
out thyroid  tissue;  (2)  administration  of  meat  to 
thyroidectomized  omnivores  or  herbivores  caused  a 
marked  increase  in  all  symptoms.  The  importance  of 
the  relation  of  the  meat  diet,  detoxication  and  thyroid 
disease  receives  considerable  confirmation  from  the  fact 
that  among  the  1,860  mammalian  postmortems  thyroid 
disease  occurred  in  2.6  per  cent,  of  all  mammals,  94.9  per 
cent,  of  which  were  found  in  flesh-eating  varieties.] 
Wells (32)  suggested  that  possibly  this  could  be  inter- 
preted as  an  indication  that  toxic  materials  found  in  the 
meat  in  the  intestinal  tract  were,  under  normal  conditions, 
detoxicated  by  the  thyroid.  Against  a  local  neutraliza- 
tion, however,  is  the  improvement  following  the 
administration  of  dried  thyroid  substance.  The  function 
is  either  neutralization  of  toxic  substances  or  the  stimu- 
lating action  on  intracellular  metabolism,  both  of  which 
might  be  called  into  play  by  an  excessive  protein  diet. 

(32)    Chemical  Pathology,  Philadelphia,,  1918. 


452   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

The  Carnivorous  Diet. 

The  pathology  of  the  more  prominent  diseases 
developed  in  carnivores  points  at  least  to  diet  as  a  pre- 
disposing or  determining  factor.  This  diet  is  very  high 
in  a  distinctly  putrefactive  protein  and  yields  products, 
chemical  and  bacterial,  which  are  toxic  and  which  give 
rise  to  acute  or  more  often  chronic  diseases  of  the  ali- 
mentary tract  and  its  adnexa.  By  reason  of  the  amount 
ingested,  excessive  because  of  lack  of  exercise,  there  is 
a  severe  tax  on  the  storage  organs  and  on  the  detoxica- 
ting  glands,  as  the  liver  and  thyroid.  The  constant 
absorption  of  these  toxic  substances  gives  rise  to  chronic 
degenerative  or  fibrotic  changes  in  the  organs  through 
which  they  pass :  liver,  kidneys,  arteries,  heart.  In  birds 
the  degenerative  diseases  are  even  more  marked  than  in 
mammals  on  the  same  diet.  The  ultimate  fault  of  this 
diet,  especially  for  mammals  and  birds  with  restricted 
activity,  lies  in  the  production  of  toxic  bodies,  produced 
either  in  the  incomplete  degradation  or  oxidation  of  the 
protein  molecule  or  as  the  result  of  bacterial  action  on  the 
protein  molecule,  a  poisonous  quality  which  is  probably 
enhanced  by  the  chemical  changes  occurring  while  the 
digested  protein  is  passing  through  the  intestinal  mucosa. 
Garden  conditions  are  such  that  these  factors  are  almost 
un surmountable  unless  the  substitution  of  vegetable  pro- 
tein could  be  accomplished.  Failure  is  often  caused  by 
limited  feeding  to  carnivores  of  muscle  and  bones, 
whereas  they  should  be  supplied  with  glandular  organs 
and  blood. 

The  Herbivorous  Diet. 

Herbivorous  diet  must  be  divided  into  two  groups, 
(1)  that  composed  of  succulent  vegetables,  and  (2)  of 
grasses,  grains  and  seeds.  In  the  first  group  there  is  an 
apparent  variation  in  the  results  found  in  mammals  and 
birds.  In  both  there  is  a  marked  decrease  in  the  chronic 
degenerative  pathology.    In  both,  acute  gastritis  is  more 


THE  RELATION  OF  DIET  TO  DISEASE       453 

prominent,  far  outstripping  the  incidence  of  this  condi- 
tion in  other  classes. 

This  diet  yields  a  large  and  quickly  available  amount  of 
carbohydrate  which  in  conjunction  with  the  moisture,  heat 
and  bacteria  which  are  unavoidably  associated  mth  raw 
vegetables,  makes  an  ideal  situation  for  infection.  These 
foods  carry  many  saprophytic  bacteria,  moulds,  etc.  In 
birds  the  conditions  are  aggravated  by  the  injuries  that 
may  occur  from  the  sharp  objects  picked  up  with  the 
gravel.  The  incidence  of  acute  infection  is  higher  among 
birds  than  among  mammals  of  this  group,  and  often  there 
is  involvement  of  the  whole  tract.  The  explanation  of  the 
other  pathological  findings  occurring  among  birds  must 
be  found  in  the  frequently  repeated  low  grade  infections 
which  result  finally  in  the  production  of  chronic  lesions 
in  the  digestive  tract,  liver,  pancreas  and  kidney.  Toxins 
as  an  etiological  factor  cannot  be  altogether  excluded,  but 
as  a  rule  they  are  not  important  because  the  by-products 
of  vegetables  are  distinctly  less  toxic  than  those  derived 
from  animal  sources.  Arteriosclerosis  is  much  less  fre- 
quent and  less  intensive  in  herbivorous  birds  than  among 
the  caiTiivorous,  probably  because  of  differences  in  the 
concentration  and  character  of  toxins  in  the  two  groups. 

Soft  Herbivorous  Diet. 

The  diet  of  succulent  vegetables  is  composed  of  tubers, 
edible  roots  and  leaves.  The  tubers  and  edible  roots  are 
high  in  water  and  carbohydrate  and  poor  in  the  amount 
and  quality  of  the  protein,  most  of  which  is  not  even  a  true 
protein  but  a  mixture  of  aminoacids.  The  leaves,  on  the 
contrary,  are  rich  in  organic  ash,  especially  calcium, 
sodium,  chlorine,  and  fat  soluble  A  vitamine,  and  as  a  rule 
contain  a  good  quality  of  protein.  They  often,  however, 
contain  injurious  substances.  This  diet,  wliile  measurably 
less  nutritious  than  that  of  the  carnivores,  can  satisfac- 
torily nourish  many  animals  with  an  extensive  intestinal 
tract  during  growth  and  even  throughout  their  entire  life. 


454  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

but  proves  entirely  inadequate  when  fed  to  an  omniv- 
orous tract. 

Seed  Diet. 

Closely  allied  in  general  character  to  the  diet  of  suc- 
culent vegetables  are  the  seed  diets,  eaten  only  by  birds 
and  having  no  parallel  among  mammalian  foods.  All 
seeds,  in  contradistinction  to  tubers,  contain  true  proteins 
which,  however,  are  of  poor  quality  because  of  the  defi- 
ciencies in  the  aminoacid  content.  They  are  as  a  rule  low 
in  the  fat  vitamines  and  in  the  amount  of  calcium,  sodium 
and  chlorine  carried.  In  three  pathological  conditions 
only  do  these  birds  show  any  oversusceptibility :  (1)  Sore 
eyes,  (2)  acute  enteritis,  (3)  osteomalacia.  Sore  eyes 
were  frequently  noted  in  this  group.  The  lesions  were 
very  like  those  described  in  animals  deprived  of  the  fat 
vitamine,  which  was  present  in  this  food  in  very  small 
amounts  or  entirely  absent,  thus  giving  a  very  plausible 
explanation  of  this  condition,  especially  as  in  some  of  the 
cases  no  other  cause  could  be  found.  Gastric  disease  of 
any  type  is  rare  in  this  group  because  the  food  at  the 
gastric  stage  is  highly  resistant  to  bacterial  action.  In 
the  duodenum,  however,  the  conditions  are  early  changed 
because  the  bacteria  carried  with  the  food  through  the 
stomach  become  active  in  the  presence  of  available  carbo- 
hydrate and  protein  decomposition  products. 

Osteomalacia  is  confined  almost  as  exclusively  to  the 
seed-eating  birds  as  it  was  to  the  omnivorous  mammals, 
and  it  is  also  associated  with  the  same  deficiencies,  cal- 
cium and  phosphorus  (cf.  Tables  19  and  20).  It  is  also 
interesting  to  note  that  these  two  diets,  the  omnivorous 
and  seeds,  yield  the  greatest  number  of  cases  of  tubercu- 
losis. Mammals  showed  32.6  per  cent.,  as  against  5.8  per 
cent,  in  all  the  other  dietary  groups,  an  observation  which 
becomes  more  striking  when  man  is  added  to  the  omnivo- 
rous group.  Seed-eating  birds  showed  17.2  per  cent.,  as 
against  6.4  per  cent,  in  other  groups.   In  both  diets  the 


THE  RELATION  OF  DIET  TO  DISEASE       455 

fat,  fat-vitamine  and  inorganic  salts,  especially  the 
calcium,  are  deficient  in  amount.  In  the  wild,  birds  vary 
their  diet  of  seeds  with  insects,  worms,  soft  fruits  and  the 
tender  shoots  of  plants,  and  at  the  same  time  they  increase 
their  inorganic  intake  by  the  minerals  picked  up  with  the 
gravel  and  from  the  water  which  has  penetrated  the  soil. ) 

Geain  and  Grass  Diet. 

The  hay-eating  animals  constitute  a  large  and  well 
studied  group — indluding  practically  all  the  domestic 
varieties.  Table  19  shows  that  these  animals  yield  the 
greatest  number  of  cases  of  malnutrition,  food  poisoning, 
acute  pancreatitis,  acute  degenerative  conditions  of  liver 
and  myocardium. 

Recent  literature  describes  many  cases  of  osteo- 
malacia, especially  among  horses  and  cows,  in  the  famine 
districts  of  Europe.  In  our  collection  of  1,860  post- 
mortems only  one  case  was  found,  that  of  an  Isabelline 
gazelle  {Gazella  isahella) ,  a  hay-eating  animal,  and  in  this 
case  it  was  secondary  to  infection. 

Arthritis,  occurring  in  3.4  per  cent,  of  all  the  autop- 
sies, was  almost  entirely  confined  to  the  hay-eating 
animals.  The  literature  describes  many  cases  of  arthri- 
tis almost  entirely  confined  to  ungulates,  of  which 
many  were  associated  with  calving  and  subsequent  infec- 
tion. Bacteriological  researches  have  found  it  most  often 
associated  Avith  streptococci,  staphylococci,  or  Bact.  per- 
fringens,  organisms  that  require  a  certain  amount  of 
carbohydrate  for  their  proper  development.  The  relation 
of  diet  to  this  condition  probably  lies  only  in  the  fact  that 
it  provides  an  excessive  carbohydrate  substrate  suitable 
for  the  optimum  development  of  these  organisms.  Folin 
and  Bergland,  noting  glycoresis  in  Herbivora,  thought  that 
it  represented  the  absorption  and  excretion  of  unusable 
carbohydrate,  present  in  grains,  vegetables,  fruits,  etc., 
and  that  it  was  sharply  separated  from  the  main  carbo- 
hydrate metabolism.    These  products  were  absorbed  from 


456   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

the  blood  exactly  as  they  were  ingested  like  lactose, 
dextrose,  etc.,  are  absorbed,  but  do  not  enter  into  the 
economy  although  they  might  cause  disorders,  especially 
forms  of  arthritis. 

The  grain  foods  are  composed  largely  of  carbo- 
hydrates (principally  in  the  form  of  cellulose  and  starch) 
small  amounts  of  protein  and  little  or  no  fat.  They  have 
a  very  low  nutritive  index  so  that  large  amounts  must  be 
consumed  to  supply  adequate  calories.  This  food  is 
constantly  present,  and  during  the  enforced  idleness  of 
capti\4ty  is  almost  continuouly  eaten.  Despite  these  facts, 
however,  malnutrition  is  present  in  2.2  per  cent,  of  the 
animals  on  this  food.  Associated  with  the  plentiful  food 
and  lack  of  exercise  are  overeating  and  pica.  Overfilled 
stomachs  occurred  thirty-four  times.  They  were  limited 
to  these  mammals  and  to  the  seed-eating  birds  whose 
environmental  conditions  are  practically  the  same.  Pica 
or  excessive  appetite  for  abnormal  food,  is  also  more  fre- 
quent in  these  groups,  but  is  usually  associated  with 
badly  balanced  diets,  and  thus  represents  an  effort  on  the 
part  of  the  animal  to  supply  its  own  deficiency.  It  is 
present  in  osteomalacic  monkeys  and  has  been  reported 
in  cattle  from  regions  where  osteomalacia  is  common  and 
following  crop  failures  where  the  rations  are  restricted. 
In  cattle  it  very  often  accompanies  food  poisoning,  espe- 
cially that  produced  by  ingestion  of  peat  hay. 

Disturbance  of  the  alimentary  tract  and  its  adnexa 
occurs  in  two  forms:  (1)  Infection  which  is  quite  com- 
mon and  involves  the  duodenum,  pancreas  and  liver,  and 
(2)  toxic.  Compared  with  other  diets  alimentary  dis- 
orders are  not  frequent  among  grain  feeders,  despite 
the  ease  with  which  grass  foods  ferment  and  the  great 
variety  of  organisms  found  in  them  such  as  moulds 
(aspergillus),  Bact.  coli,  paratyphosus,  enteritidis, 
suipestifer,  oidium  lactis,  etc.  Few  bacteria  can 
attack  whole  protein,  cellulose  or  starch,  and  the  decompo- 
sition products,  peptone,  glucose,  etc.,  are  not  available 


THE  RELATION  OF  DIET  TO  DISEASE       457 

in  any  quantity  until  the  lower  stomacli  and  duodenum 
are  reached.  The  inflammation  of  the  alimentary  tracts 
of  these  animals  is  confined  to  the  fourth  stomach  and 
duodenum,  with,  in  many  cases,  extension  to  liver 
and  pancreas. 

Acute  and  chronic  degenerative  changes  occur  very 
frequently,  and  as  a  rule  are  the  result  primarily  of 
absorbed  toxins.  After  ingestion  of  new  hay  this  often 
appears.  The  toxic  substance  probably  is  a  terpinol  ester, 
cumarin,  which  is  produced  by  an  enzyme  in  the  cut  grass. 

The  result  is  a  gastroenteritis  with  jaundice,  thirst  and 
marked  flatulence.  It  is  very  probable  that  many  of  the 
gastrointesinal  and  degenerative  lesions  are  the  result 
of  the  combined  action  of  toxin  and  bacteria. 

Food  Poisoning. 

;^  Food  poisoning  occurs  in  all  diets,  but  especially 
among  the  grass-eating  mammals.  To-day  under  the 
general  heading  of  food  poisoning  are  included  those 
cases  due  to  (1)  some  injurious  substance  inherent  in  the 
food  itself,  true  food  poisoning,  (2)  those  due  to  toxic 
substances  liberated  or  produced  in  food  contaminated  by 
parasites  or  bacteria,  (3)  those  due  to  bacteria  that  are 
carried  by  food  and  develop  into  true  infection  after 
ingestion.  Most  of  the  cases  of  meat  poisoning  described 
in  literature  undoubtedly  belong  to  this  third  class,  i.e., 
flesh  is  infected  during  the  life  of  the  animal  or  during 
its  preparation  for  food  and  the  virus  develops  in  the 
host  after  ingestion.  A  fourth  and  more  rare  class  of 
food  poisoning  is  due  to  the  condition  of  the  individual 
consuming  the  food — ^protein  sensitization. 

Injurious  constituents  of  normal  flesh  foods  are  very 
micommon.  There  are  a  few  poisonous  fish,  notably  the 
balloon,  puffer,  and  Fuga  fish  of  Japan,  which  when 
eaten  give  rise  to  cholera-like  conditions  ending  in  con- 
vulsions and  paralysis.  A  marked  intoxication  has  been 
described  in  dogs  which  have  fed  upon  the  Greenland 

■'       30 


458   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

shark.  Some  fish  are  poisonous  at  certain  periods  as 
spawaiing  season,  the  poison  then  being  confined  to  the  roe. 
Still  others  are  harmless  miless  rendered  toxic  by  some 
injurious  food.  This  poisoning  of  muscle  meats  is  seen 
in  quail  and  partridges  fed  on  mountain  laurel,  in  some 
fish  after  consuming  certain  marine  plants,  and  in  cattle 
poisoned  by  amanita. 

The  most  common  sources  of  poisoning  are  spoiled 
meat  and  flesh  of  diseased  animals,  both  of  which  are 
serious  factors  in  the  production  of  the  gastrointestinal 
disorders  of  omnivores  and  carnivores.  Practically  all 
the  reports  of  meat  poisoning  from  the  literature  have 
been  traced  to  the  use  of  raw  or  insuflficiently  cooked 
flesh,  and  have  yielded  on  bacteriologic  examination  Bad. 
paratypliosus,  Bad.  enteritidis,  Bad.  suipestifer,  Bad. 
coll,  or  Bad.  proteus. 

The  bacteria  may  produce  toxin  in  the  food  previous 
to  ingestion  causing  in  the  host  only  a  severe  intoxication. 
This  is  the  situation  developed  after  eating  sturgeon 
infected  with  Bad.  piscidus  agilis,  an  organism  which 
manufactures  a  highly  poisonous  alkaloid.  A  similar 
intoxication  follows  the  ingestion  of  potatoes  infected 
with  Bad.  proteus  or  containing  the  poisonous  alkaloid, 
solanin,  which  is  produced  in  diseased  and  sprouting 
potatoes.  Other  examples  of  this  are  (1)  ergotism — due 
to  an  infection  of  rye  and  wild  grasses  with  Claviceps 
purpurea  which  produces  three  poisonous  bodies,  ergo- 
tinic  acid,  which  is  not  poisonous  when  taken  into  the 
stomach,  sphacetinic  acid  and  cornutin  which  act  on  the 
nervous  system,  brain,  cord,  vagus  and  vasomotor  centre 
giving  rise  to  toxic  polyneuritis,  and  (2)  favus,  an  acute 
febrile  anemia  with  jaundice  and  hemoglobinuria  prob- 
ably due  to  a  bacterial  infection  or  fungus  growth  of  the 
bean.  Infected  food  may  also  produce  soluble  heat- 
resisting  toxins  that  produce  immediate  symptoms  and 
increase  the  animal's  susceptibility  to  infection.  This  is 
the  more  common  finding  in  cases  of  poisoning  with  milk 


THE  RELATION  OF  DIET  TO  DISEASE       459 

and  milk  products.  Non-pathogenic  saprophytes  carried 
in  milk  produce  (1)  a  poison  closely  allied  to  tyrotoxicon, 
(2)  a  toxalbumin  which  in  itself  causes  serious  disturb- 
ances. Botulism,  also  probably  of  this  group,  is  a  disease 
initiated  by  a  toxin  elaborated  by  Bad.  hotulinus  acting 
on  a  protein.  There  is,  however,  some  evidence  that  Bad. 
hotulinus  can  also  establish  a  real  infection. 

The  toxemias  from  food  infected  with  bacteria  may  not 
occur  until  the  food  is  ingested  or  the  bacteria  implanted. 
This  result  occurs  in  infections  with  Bad.  bovis  morhif- 
icans,  Gartner's  bacillus,  etc.,  or  after  the  feeding  of 
meat  from  animals  infected  with  Bad.  paratyphosus 
and  enteritidis. 

The  plant  poisons  are  more  frequently  due  to  inherent 
injurious  substances,  although  even  among  them,  bac- 
terial and  fungus  diseases  play  an  important  role. 
Among  the  16,673  plants  indigenous  to  North  America, 
almost  500  are  more  or  less  poisonous  and  about  30 
are  of  great  economic  importance.  The  toxic  factor  may 
be  confined  to  the  leaf,  seed  or  root,  but  more  often  it  is 
associated  with  all  parts  of  the  plant.  Through  the  efforts 
of  the  Department  of  Agriculture  a  more  or  less  complete 
list  of  the  plants  implicated  in  the  poisoning  of  stock  has 
been  compiled.  This  list  includes  the  following :  Amanita 
muscaria;  A.  phalloides;  Veratrum  viride;  Phytolacca 
decandra;  Agrostemma  githago;  Delphinium,  25  varie- 
ties; Astragalus  mollissimus;  Aragallus  lamhertii; 
Crotalaria  sagittalis;  Euphorbia  lathyris;  E.  marginata; 
Rhus  radicans;  R.  diver siloba;  R.  vernix;  Aesculus  pavia; 
A.  hippocastanum;  A.  glabra;  A.  Calif ornica;  Cicuta 
maculata;  C.  vagans;  Conium  maculatum ;  Kalmia  lati- 
folia;  K.  augustifolia;  Leucothoe  catesbaei;  Rhododen- 
dron maximum;  Pieris  mariana;  Datura  stramonium; 
Solanum  nigrum;  S.  dulcamara;  Helenium  autumnale; 
Asclepias  pumila;  A.  verticullata;  A.  galoides;  A.  mexi- 
cana;  A.  eriocarpa;  A.  speciosa;  A.  fremonti;  Eupa- 
torium  agertoides;  E.  urticar folium;  Isocoma  wrightii; 


460  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Dauhentonia  longifolia;  Senecio  jacohia  hurchelli  latifo- 
Zw5.(33)  Some  of  these  as  the  Amanita  are  only  occasional 
sources  of  disaster,  but  as  they  frequently  involve  man 
they  are  important.  The  Amanita  muscaria  symptoms 
appear  very  soon  after  eating  the  fungus  and  consist  of 
a  deepening  stupor.  A.  phalloides,  on  the  contrary,  starts 
with  severe  abdominal  pain,  cramps,  discharges  of  blood 
and  mucus  and  later  convulsions.  The  meat  of  animals 
dying  from  fungus  poisoning  is  distinctly  poisonous. 
This  transfer  of  poison  to  the  muscles  of  the  animal  par- 
taking of  these  plants  occurs  also  in  poisoning 
with  Kalmia. 

The  other  plants  of  this  list  are  closely  associated 
with  the  grass  foods  and  are  consumed  usually  when  the 
food  on  a  range  is  scarce.  Some  groups  as  the  Asclepias 
contain  a  distinct  neurotoxin  and  give  rise  to  a  condition 
known  as  trembles  or  staggers.  It  affects  mostly  cows 
and  sheep,  causing  staggering,  trembling  gait,  bloating 
and  salivation  and  death  with  convulsions.  There  is 
marked  congestion  of  alimentary  tract,  liver  and  kidney. 
In  the  cerebrospinal  axis  there  are  marked  changes  in  the 
nerve  cells  of  the  medulla  and  spinal  cord.  The  Purkinje 
cells  show  the  effect  of  extreme  fatigue.  Other  plants 
causing  stiffness  or  weakness  of  the  extremities,  show 
on  microscopic  examination  no  definite  lesions  in  the 
cerebrospinal  axis.  Loco  weed — Astragalus  mollissimus 
and  Aragallus  lamhertii — causes  maniacal  disturbances 
but  no  gross  lesions.  This  weed  in  Colorado  costs  the 
state  enormous  amounts  of  money  yearly. 

Helenium  poisons  domestic  animals  by  means  of  a 
toxic  glucoside,  dugaldin,  which  produces  stiffness,  sali- 
vation and  nausea  mth  mild  depression  (''  spemng 
sickness  ") .  The  alimentary  tract  shows  severe  inflamma- 
tion of  the  rumen  and  reticulum  which  may  at  times  be 
hemorrhagic.    The  Hver  usually  presents  an  interstitial 

(33)  These  botanical  names  are  taken  from  Chestnut's  Poisonous 
Plants  of  America. 


THE  RELATION  OF  DIET  TO  DISEASE       461 

hepatitis.  This  toxin  is  decidedly  hemolytic.  The  effects 
of  this  plant  are  always  permanent,  total  recovery  being 
very  rare. 

The  larkspur  (25  different  varieties),  on  the  contrary, 
shows  prompt  recovery  after  treatment,  but  no  establish- 
ment of  toleration.  These  plants  give  rise  to  nausea, 
vomiting  and  great  agitation  and  destroy  many  animals 
yearly.  The  poisons  are  included  in  four  alkaloids,  all 
spinal  cord  depressants  resembling  aconite  in  general 
character. 

These  poisonous  plants  all  produce  more  or  less  gas- 
trointestinal inflammation  and  practically  all  are 
destructive  in  their  action  on  the  liver,  pancreas  and 
kidney.  It  is  impossible  to  form  even  approximate  esti- 
mates of  the  damage  done  by  them  because  of  the  general 
ignorance  of  the  subject.  The  Division  of  Botany  has 
been  collecting  for  the  past  few  years  specific  information 
concerning  these  plants,  but  the  individual  plants  are  not 
equally  poisonous,  and  all  animals  do  not  show  the  same 
susceptibility  to  the  poison.  Veratrum  viride,  for 
instance,  is  eaten  with  relish  by  sheep  and  elk  and  is 
decidedly  toxic  for  the  horse.  In  many  the  toxic  factor 
has  not  been  isolated.  Some,  as  Euphorbia,  are  poisonous 
only  when  fed  in  honey  derived  from  its  flowers. 

The  influence  of  diet  on  the  general  health  of  animals 
is  very  far  reaching  and  very  inclusive.  Metabolic  dis- 
turbances are  undoubtedly  at  times  the  result  of 
unbalance — deficiencies  on  the  one  side,  excesses  on  the 
other,  at  times  are  probably  much  more  the  results  of 
bacterial  invasions  aided  and  abetted  by  the  food  admin- 
istered, at  still  other  times  are  poisonous  either  in  their 
own  content  or  from  the  degradation  products  resulting 
from  digestion  or  bacterial  decomposition. 


SECTION  XVI 

NEOPLASMS 

The  occurrence  of  true  neoplasms  in  domesticated 
animals  has  always  been  well  known  and  thoroughly 
studied  while  for  beasts  in  the  wild  the  data  has  been 
fragmentary.  That  tumors  exist  in  natural  environment 
has  been  accepted  upon  the  testimony  of  hunters  but 
there  is  an  impression,  and  nothing  more,  of  their 
extreme  scarcity  probably  because  only  younger  vig- 
orous animals  come  to  the  attention  of  the  sportsman 
or  collector.  This  matter  will  of  course  not  be 
settled  until  some  natural  historian  with  a  knowledge 
of  pathology,  makes  a  survey  of  a  large  number  of  speci- 
mens taken  during  a  collecting  expedition.  Observations 
in  menageries  are  valuable  to  the  extent  that  they  show 
what  tumors  may  occur,  the  orders  most  commonly 
affected  and  the  incidence  under  captive  conditions.  It 
is  unfortunate  that  too  seldom  do  we  know  the  history  of 
our  specimens  in  regard  to  the  age,  manner  of  capture 
or  breeding,  data  which  if  at  our  disposal  would  permit 
of  a  very  fair  idea  of  the  probable  incidence  in  the  wild 
and  of  the  effect  of  captivity.  Some  observations  in  this 
direction  are  however  possible  by  using  the  figures  of 
known  captivity  and  breeding. 

The  facts  gleaned  from  a  study  of  neoplasms  under 
captive  conditions  may  be  of  interest  to  the  experimental 
pathologist,  especially  when  considering  the  relation  of 
the  origin  from  the  embryological  layers.  I  have  tabu- 
lated this  with  great  care,  using  Jordan's  (1)  table  for 
the  source  of  the  various  tissues,  and  further  have  studied 
the  destination  of  metastatic  emboli  in  terms  of 
the  blastoderm. 

The  following  observations  are  based  entirely  upon 
our  own  data  for  while  it  might  be  valuable  to  include  the 
cases  in  the  literature  their  descriptions  are  often  so 

(1)    Textbook  of  Histology,  1920. 
462 


NEOPLASMS  463 

meagre  that  they  would  not  combine  readily  with  our 
records.  Plimmer,  Seligmann,  and  Murray  have  pub- 
lished in  the  Proceedings  of  the  London  Zoological 
Society  since  1903,  their  annual  report  of  the  pathological 
service  in  which  they  have  recorded  very  many  interest- 
ing tumors.  So  too  from  time  to  time  Harlow  Brooks 
and  W.  R.  Blair  in  the  Annual  Report  of  the  New  York 
Zoological  Park,  have  presented  cases  occurring  in  their 
ser\dce.  Joest  (2)  discusses  tumors  in  the  lower  animals 
in  a  broad  way  and  analyzes  their  incidence  and 
characters.  C.  Y.  White  and  I  (3)  have  already  published 
articles  on  this  subject.  Numerous  single  references  may 
be  found  in  the  Jahresher.  der  Veterindr-Medicin. 

In  so  far  as  the  incidence  of  tumors  in  wild  animals  is 
concerned  this  literature  can  scarcely  give  an  adequate 
measurement  but  it  would  seem  that  they  are  less  than 
in  domestic  varieties.  Exact  figures  for  the  occurrence 
of  tumors  in  the  latter  seem  not  available  in  the  literature, 
but  one  can  find  that  in  the  Prussian  army  horses  about 
one  hundred  are  obser\^ed  each  year.  In  our  5,365  speci- 
mens collected  during  nineteen  years,  94  tumors  in  92 
animals  have  been  found,  1.7  per  cent,  or  about  one  in 
every  sixty  specimens,  not  at  all  a  low  figure.  If  one  were 
to  include  all  fibromata  of  the  feet  and  the  blood  collec- 
tions to  which  the  name  angioma  might  be  applied,  this 
incidence  would  be  greater;  they  are  excluded  because 
few  in  number  and  vag-ue  in  history;  only  one  true 
angioma  was  seen. 

The  gross  and  microscopical  appearances  of  tumors  in 
the  lower  animals  are  essentially  the  same  as  one 
encounters  in  human  beings  or  at  least  it  is  possible  to 
apply  the  pathological  nomenclature  used  in  human 
medicine  to  all  the  neoplasms  we  have  discovered.  There 
is  appended  a  list  of  all  the  animals  and  their  tumors,  a 
table  of  zoological  orders,  tumors  and  organs  (Table  21) 
and  an  analytical  table  of  the  histological  data.  (Table  22) 

(2)  Zeitch.  fiir  Krebsforsch,  Vol.  15,  p.  1. 

(3)  Proceed.  Phila.  Path.  Soc,  1910,  and  Journal  of  Pathology  and 
Bacteriology,  Vol.  XVII,  1912. 


464   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 


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468  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Incidence  of  Tumors. 

Examination  of  the  table,  (21)  from  the  standpoint 
of  differential  percentage  reveals  that  mammals  have  48 
tumors  giving  an  incidence  of  2.58  per  cent,  whereas  birds 
have  44  newgrowths  equivalent  to  1.23  per  cent.  Were 
it  not  for  the  high  figures  for  one  single  variety  of  bird 
(Undulated  Grass  Parrakeet)  this  value  for  Aves  would 
be  still  lower.  At  all  events  our  figures  would  indicate 
that  the  mammal  is  at  least  tmce  as  productive  of  neo- 
plasms as  is  the  bird.  In  our  material  the  latter  class  has 
had  a  better  chance  than  Mammalia  to  show  its 
susceptibility  since  there  have  been  nearly  t-\vice  as 
many  autopsies. 

Within  the  classes  the  comparative  figures  have  less 
value  because  of  the  smaller  and  varying  numbers.  Such 
high  percentages  as  are  shown  by  the  elephants  and 
armadillos  cannot  be  taken  as  indicators  for  their  orders 
since  too  few  specimens  were  examined.  Judging  by 
orders  with  more  than  one  hundred  autopsies  the  rodents 
stand  at  the  head  of  the  list  followed  by  the  marsupials 
and  carnivores.  It  is  interesting  that  the  animal  nearest 
to  man,  the  monkey,  and  with  greatest  number  of  autop- 
sies in  its  zoological  class  has  the  lowest  tumor  incidence. 
Psittaci  lead  the  avian  orders,  followed  by  the  Fulicariae, 
but  as  there  are  but  thirty-five  autopsies  upon  these,  the 
second  place  rightly  belongs  to  the  Anseres.  All  the 
principal  orders  are  represented  but  the  only  one  of 
importance  is  the  leader.  The  Psittaci  are  very  prone  to 
have  tumors  in  the  renal  area,  sometimes  of  the  kidney, 
at  others  of  the  adrenal  and  occasionally  of  the  sex 
glands.  Some  remarks  have  already  been  given  to  this 
matter  in  the  sections  devoted  to  the  Iddney  and  genitalia 
but  it  mil  be  discussed  again  under  tumor  morphology. 

Among  these  ninety-two  animals,  one  bore  multiple 
tumors,  a  Jaguar  {Felis  onca)  with  adenomata  of  the 
liver  and  uterus  and  angiomata  of  the  mesentery.    Care- 


NEOPLASMS  469 

ful  study  failed  to  reveal  any  parasitism  as  the  cause  of 
the  growths  and  since  the  first  two  were  of  slightly  vary- 
ing structure  it  is  not  believed  that  one  is  a  metastasis 
from  the  other. 

The  sex  incidence  stands  in  direct  relation  to  the  pro- 
portion of  total  males  and  females  posted  or  in  other 
words  it  is  the  same  for  the  two.  The  figures  might  be 
somewhat  affected  were  the  gender  of  all  the  parrakeets 
available  but  the  tumors  growing  in  the  upper  renal  area 
frequently  destroy  the  sex  gland. 

Definite  statements  concerning  the  importance  of 
breeding  in  the  causation  of  neoplasms  cannot  be  made 
since  we  cannot  quote  figures  for  the  percentages  of  wild- 
and  captive-born  of  our  entire  autopsy  list.  The  data 
are  confused  by  scanty  information  concerning  the 
twenty-six  parrots,  the  history  of  which  is  vague  and  I 
am  perhaps  too  severe  in  accrediting  the  birth  of  sixteen 
of  them  to  capti\T.ty.  Tliis  was  done  because  of  a  lack  of 
exact  information  concerning  these  specimens  and,  be- 
cause their  variety  is  lmo\\Ti  to  breed  when  captive  by  the 
residents  of  their  habitat(4),  the  distribution  into  wild- 
and  captive-born  is  based  upon  what  information  we  have. 
If  the  order  Psittaci  be  subtracted  entirely,  it  leaves  a 
total  of  62  tumor-bearing  animals  of  kno^vn  breeding,  49 
of  which  were  born  in  the  wild,  thirteen  in  captivity,  a 
fact  which  strengthens  the  thought  that  unnatural  breed- 
ing increases  the  chance  of  neoplasms. 

The  kno^vn  length  of  capti^ity  has  also  a  direct  bearing 
on  this  point.  The  figures'  given  in  the  columns  "  known 
captivity  "  and  ^'  average  for  tumor  bearers  "  were 
compared  with  figures  obtained  by  averaging  the  lives  of 
fifty  others  (when  possible)  of  the  same  order  or  of  at 
least  three  times  as  many  as  bore  tumors.  Animals 
dying  from  injury  were  excluded.  With  one  exception 
the  average  for  '^  tumor  bearers  "  exceeded  that  for 
"non-tumor  bearers";  the  exception,  the  Ungulata,  had 

(4)    See  Gould's  Birds,  Vol.  II,  p.  83. 


((' 


470  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

the  same  average  for  both  groups.  It  seems  then 
that  tumors  occur  in  animals  in  captivity  longer 
than  the  average  for  their  order,  or  in  specimens 
that  have  the  power  to  live  under  confined  conditions  until 
neoplasms  develop.  In  this  respect  I  recall  the  state- 
ments made  by  Harlow  Brooks  (5)  that  tumors  will  prob- 
ably be  found  more  commonly  in  animals  when  they  live 
in  a  manner  comparable  to  that  of  urban  man  and  that 
racial  degeneracy  will  favor  their  development.  There 
is  adduced  here  perhaps  the  first  definite  evidence  that 
long  captivity  allows  tumor  tendency  to  express  itself 
but  it  does  not  prove  that  confinement  increases  tumors. 
Nor  does  the  expectation  of  life,  average  or  potential, 
stand  in  any  direct  and  definite  relationship  to  the  fre- 
quency of  neoplasms.  The  only  clear  case  of  long  life 
and  high  tumor  incidence  is  to  be  found  in  Parrots ;  we  feel 
however  that  some  miknown  factor  increases  tumors  of 
the  renal-adrenal  region  in  these  birds  and  that  unquali- 
fied statements  about  age  and  tumor  growth  are  not  per- 
missible. Since  tumors  grow  in  many  wild-born 
specimens,  a  high  percentage  of  which  become  known  in 
the  first  few  years  of  captivity,  is  it  not  highly  probable 
that  tumors  are  reasonably  common  in  the  wild  and  that 
we  do  not  observe  enough  purely  natural  specimens  to 
assume  an  immunity  on  the  part  of  free  li\dng  beasts. 

One  of  the  undesirable  features  of  captive  breeding  is 
consanguinity  of  parents  and  there  is  good  reason  to 
believe  that  tumor  susceptibility  can  be  bred  into  or  out 
of  a  line  of  animals  by  mating  tumor  bearers  and  non- 
I  tumor  bearers,  the  tendency  following  the  rules  of 
Mendelian  inheritance  (Slye).  Is  there  any  proof  that 
inbreeding  does  not  occur  in  the  wild  and  if  it  do,  it  is 
perfectly  possible  that  tumor  tendency  may  be  trans- 
mitted as  a  dominant  character ;  the  effect  of  artificial  or 
intentional  inbreeding  in  captivity  would  only  offer  an 
opportunity  for  a  summation  of  these  influences. 

(5)   Am.  Jour.  Med.  Soc,  1907,  133-769. 


NEOPLASMS  471 

If  injury  and  animal  parasitism  have  any  importance 
in  neoplasmata  then  this  opportunity  certainly  occurs 
under  natural  conditions,  Fibiger  observed  gastric 
tumors  in  rats  arising  under  the  influence  of  nematodes 
while  Slye  and  Wells  report  facial  neoplasms  in  mice 
apparently  arising  at  points  of  old  injuries.  It  seems  to 
me  that  we  have  no  right  to  assume  an  immunity  of  wild 
animals,  in  their  native  environment,  to  tumors;  the 
incidence  is  another  matter  but  it  may  be  considerable. 

It  was  thought  possible  that  there  might  be  some  light 
shed  upon  the  matter  by  an  analysis  of  our  sarcomatous 
and  epitheliomatous  tumors  in  wild-  and  captive-bom 
animals.  In  our  second  paper  (6)  upon  this  subject  I 
ventured  the  statement  that  sarcomatous  growths 
occurred  more  frequently  in  captive-born,  epithelioma- 
tous in  wild-born  specimens.  Greater  data  have  not  borne 
out  this  conclusion  and  information  was  sought  as  to  the 
embryonal  derivation  of  tumor-bearing  tissue.  Analyz- 
ing the  cases  in  which  all  the  factors  could  be  obtained,  it 
seems  that  among  seven  tumors  of  captive-bred  animals, 
five  came  from  the  entoderm,  two  from  the  mesoderm, 
whereas  in  wild-bred  animals,  of  the  fifty-seven  tumors, 
five  came  from  the  ectoderm,  thirty-two  from  the 
mesoderm  and  fourteen  from  the  entoderm.  These 
figures  do  not  include  the  parrots.  The  sex  values  have 
no  significance. 

It  is  interesting  and  noteworthy,  that,  as  in  the  human 
being,  the  majority  of  the  tumors  came  from  tissues  aris- 
ing in  the  mesoderm  and  that  the  entodermic  derivatives 
received  the  largest  number  of  metastases ;  no  ectodermic 
tissues  were  sites  of  secondary  tumors.  The  visceral 
seats  of  metastases  are  probably  of  little  value  for  com- 
parison in  so  small  a  number ;  the  lung  and  liver  however 
occupy  the  prominent  places. 

Interesting  as  the  foregoing  facts  may  be,  they  do  not 
shed  light  upon  the  question  of  breeding  and  degeneracy 

(6)   Jour.  Path,  and  Bact.,  Vol.  XVII,  1912. 


472  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

in  the  causation  of  neoplasms.  Attention  is  arrested  how- 
ever by  the  paucity  of  tumors  in  derivatives  of  the  ecto- 
derm since  in  man  new  growths  are  common  in  the  breast, 
at  the  rectal  and  labial  mucocutaneous  jmictions  and  on 
the  skin.  The  immunity  of  the  ectodermic  tissues  to 
secondary  growths  is  very  distinct ;  this  holds  true  in  man. 

Special  Tumoes. 

The  diagnosis  of  fibroma  offers  the  same  difficulty  in 
the  zoological  material  as  it  does  in  man  and  even  more 
care  must  be  exercised  for  solid  tumors  in  certain  locali- 
ties. The  bird  often  presents  hard  nodular  masses  on  the 
palmar  and  lateral  aspects  of  the  feet,  sometimes  sur- 
mounted by  callosities,  to  which  the  term  fibroma  or  fibro- 
matous  corns  might  be  applied.  Section  of  some  of  these 
will  reveal  areas  of  granulation  tissue  about  points  of 
inflammation  so  that  we  have  considered  them  as  infec- 
tious or  the  result  of  incorrect  perches  and  excluded  them 
from  the  tumors.  True  fibromata  have  been  encountered 
thrice  but  in  combination  with  muscle  tissue  as  a  fibro- 
myoma  thrice  in  addition.  The  ''  fibroids  "  seen  in  the 
elephants  and  armadillo  have  already  been  described. 

The  nodular  growth  sometimes  accompanying  degen- 
erative disease  of  the  osseous  system  followed  by 
attempts  at  repair  as  discussed  under  osteitis  deformans, 
leontiasis  ossium  and  actinomycosis,  are  often  produc- 
tive of  masses  to  wliich  it  is  easy  to  apply  the  term 
osteoma.  If  one  demand  that  an  osteoma  shall  be  a  dis- 
tinct neoplastic,  localized  bony  growth  of  unnatural  or 
greatly  exaggerated  structure,  then  the  tumor  is  quite 
rare.  We  have  seen  one  growing  from  the  vertebrae  and 
clavicle  of  a  gerbille  and  a  fibro-osteoma  on  one  jaw  of  an 
Isabelline  gazelle.  The  chondromata  have  been  limited 
to  one  case,  a  unilateral  mass  growing  from  the  nasal 
cartilage  of  a  caracal. 

(  Lipomata  are  localized  collections  of  fat  consisting  of 
cells  with  greater  fat  capacity  than  normally,  sometimes 


NEOPLASMS  473 

surrounded  by  an  indefinite  capsule.     Judging  by  the  A 
observations  of  Joest  and  Johne  they  are  reasonably  \ 
common  in  horses  and  cows.    We  have  not  seen  a  single  ' 
case  in  mammals  but  eight  cases  appeared  in  the  birds. 
These  were  with  one  exception  disposed  under  the  skin 
mostly  over  the  abdomen  and  chest  and  once  under  the 
scalp.    In  a  hawk  the  tumor  grew  as  a  pehdc  mass  sur- 
rounding  the    cloaca    and    apparently    caused    decided 
obstruction  to  the  lumen.    The  lipomata  of  the  Psittaci 
usually  grow  as  pendulous  masses  on  the  abdominal  wall 
covered  by  thin,  featherless,  delicate  skin,  often  show- 
ing dilated  veins.    Upon  section  they  are  rather  rich  in 
blood   supply,   "  angiolipoma,"   but   fail   to    show   any 
angiomatous  or  solid  cellular  areas  under  the  micro- 
scope.    The  frequency  of  the  growths  in  one  variety 
(Roseate    cockatoo — Cacatua    roseicapilla)    led    to    an 
attempt  to  transplant  the  tumor.    The  plant  seemed  to 
thrive  in  the  recipient  for  a  while  but  soon  disappeared,  j 
Breeding  experiments   on  the   tumor-bearers   are   now 
under  way. 

Angiomata  of  lymph  channels  were  observed  in  the 
omentum  and  mesentery  of  a  jaguar  (Felis  onca) ;  this 
is  the  animal  with  three  apparently  separate  and  dis- 
tinct tumors.  "  The  omentum  is  normally  fatty  and 
slightly  congested.  In  its  meshes  are  myriads  of  tiny 
cysts  containing  gray  fluid.  The  main  peritoneal  area 
is  negative  but  in  the  pelvic  region  on  anterior  rectal 
wall,  in  the  superior  edge  of  the  broad  ligament  and  in 
Douglas'  pouch,  are  cysts  from  a  few  millimetres  to 
several  centimetres,  with  clear  contents.  The  microscopic 
section  of  omentum  shows  the  multiple  cysts  as  cavities  of 
varying  size,  from  that  of  an  arteriole  to  the  diameter  of 
a  two-third  lens  field.  They  are  lined  with  flat,  closely 
placed  pavement  cells  with  well  stained  but  vesicular 
nucleus.  The  septa  are  adult  connective  tissue.  No  con- 
tents or  gTanular  eosin-staining  material.  No  swollen 
cells  like  in  adenomata.  No  parasites  seen." 
31 


474  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Two  endotheliomata  have  been  found,  one  of  the  flat 
variety  with  warty  excrescences  common  on  serous  sur- 
faces, located  in  the  pleura  of  a  leopard  {Felis  nebulosa), 
and  one  of  the  nodular  variety,  growing  from  the  clavicle 
of  a  Moorhen  {Gallinula  chloropiis). 

The  sarcomata  present  their  usual  morphology 
grossly  and  minutely  and  with  the  exception  of  the  cases 
arising  from  the  pectoral  muscle  and  from  the  genital 
area  offer  little  of  interest.  Two  instances  in  the  former 
location,  observed  in  parrakeets,  presented  several  puz- 
zling features.  The  component  cells  were  spindle  in  shape, 
similar  to  a  muscle  cell  but  were  fitted  with  the  round  or 
elliptical  nuclei  of  embryonal  cells.  In  a  few  places  they 
were  exceedingly  large  and  had  shadowy  outlines  like  a 
syncytium  or  they  would  be  so  arranged  as  to  suggest  a 
glandular  structure.  The  dominant  type  of  cell  was,  how- 
ever, everywhere  the  spindle  as  it  is  seen  in  sarcoma.  The 
sarcomata  when  they  occur  in  the  genital  area  usually 
assume  the  alveolar  arrangement  and  are  of  the  round  or 
mixed  cell  variety.  Only  three  of  the  sixteen  sarcomata 
gave  metastases. 

Papillomata  of  minor  character  appear  occasionally 
on  the  skin  of  animals  as  warts,  but  only  one  instance  of 
any  greater  importance  has  been  found.  The  duodenal 
mucosa  of  an  owl  {Bubo  virginianus)  presented  a  soft 
growth  which  partly  obstructed  the  intestinal  lumen. 
Papillary  adenomata,  on  the  other  hand,  have  been 
observed  several  times,  but  since  they  have  more 
importance  as  irregular  hyperplasias  of  glandular  origin 
have  been  included  in  the  next  group.  An  interesting 
case  was  seen  in  a  baboon  {Papio  hamadryas)  in  which  a 
large  part  of  the  gastric  wall  was  the  seat  of  adenomata, 
presenting  in  addition  several  distinct  papillary  out- 
growths. ^  A  similar  picture  was  found  in  the  duodenum 
of  the  rhea  {Rhea  am  eric  ana).  ) 

The  greatest  interest  in  the  adenomata  centres  around 
these  growths  in  the  renal  area  in  parrakeets,  and  as  they 


NEOPLASMS  475 

have  much  in  common  with  all  the  glandular  tumors  of 
this  region,  a  general  discussion  of  this  subject  may  be 
introduced  here.  We  have  observed  seven  tumors  con- 
structed on  a  glandular  basis  of  renal  or  adrenal 
character.  Grossly  these  tumors  develop  as  irregular 
masses  usually  of  distinct  brown  color,  constructed  on  a 
lobular  plan,  delicate  barely  visible  septa  dividing  the 
growth.  They  seem  devoid  of  large  vessels,  a  gross 
observation  confirmed  microscopically.  There  is  no  cri- 
terion to  the  naked  eye,  which  will  distinguish  the  variety 
of  epithehal  hyperplasia  or  permit  separation  of  these 
neoplasms  from  some  sarcomata;  the  latter  are  usually 
gray  but  need  not  be  so.  Minutely  studied,  three  of  these 
tumors  proved  to  be  adenomata,  all  papillary,  one  cystic 
as  well.  Three  had  to  be  denominated  carcinoma  because 
of  their  distinct  separate  crowded  nests  and  incomplete 
acini.  The  cells  comprising  these  growths  are  comparable 
to  the  lining  elements  of  the  collecting  tubules  of  the  renal 
lobule  in  that  they  have  relatively  large  nuclei  and  a 
tendency  to  basic  staining  protoplasm.  The  adenomatous 
picture  is,  however,  more  comparable  to  the  cortex  than  to 
the  medulla.  The  remaining  tumor  was  a  hypernephroma 
of  the  usual  large  cell,  acinus-forming  type  and  seemed 
to  originate  in  the  adrenal.  None  of  these  tumors  in  the 
parrakeets  sent  out  metastases.  Other  hj^Dernephromata 
have  been  diagnosed,  to  the  number  of  six.  Upon  review 
of  their  descriptions  and  sections,  the  determinations  are 
to  be  confirmed.  However,  it  must  be  recorded  here  that 
none  of  the  three  in  mammals  gave  metastases,  while  two 
of  the  three  in  birds  did  so.  They  are  all  of  the  usual  type 
with  large  vacuolated  cells  in  glandular  groups  or  strands. 
Three  rather  interesting  examples  of  epithelioma  have 
been  observed.  The  first  and  most  important  was  a  baso- 
cellular  growth  of  the  tongue  in  a  black  bear  (Ursus 
americanus).  The  local  damage — ulceration  and  infiltra- 
tion— and  swelling  sufficient  to  interfere  with  deglutition, 
were  quite  considerable.    The  basal  cell  nests  had  pene- 


476   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

trated  deeply  into  the  muscle,  but  extension  had  taken 
place  only  to  a  single  submaxillary  gland.  A  squamous 
epithelioma  was  found  on  the  skin  of  the  thigh  of  a  Tas- 
manian  devil  {Sarcophilus  wsiniis).  The  construction 
was  somewhat  unusual  in  that  it  was  cystic  but  lined  with 
squamous  and  keratinized  plates.  It  could  not  be  decided 
that  it  originated  from  glands  like  a  trichoepithelioma; 
it  was  not  like  a  basal  cell  cancer.  No  metastases  had 
occurred.  The  third  case  was  that  of  a  tumor  mthin  the 
abdomen  of  an  Amazon  {Chrysotis  leucocephala).  It  con- 
sisted of  an  illy  defined  basement  membrane  upon  which 
were  irregular  stratified  squamous  epithelial  cells.  Upon 
the  surface  were  wavy  bands  of  horny  material,  very 
much  like  dried  and  cast-off  epithelial  scales,  except 
more  compact  and  extensive.  These  latter  seemed  to 
form  the  bulk  of  the  mass.  Beneath  the  membrane  a  few 
irregular  accumulations  of  cells  bearing  a  similarity  to 
those  on  the  surface  could  be  found,  but  they  were  prob- 
ably large  plasma  cells.  *The  epithelial  layer  dipped  down 
like  in  epithelioma.  No  pearls  or  separate  nests  were 
found.  Wliile  this  mass  was  not  localized,  it  was  doubt- 
less an  epithelioma,  and  should  be  included  in  this 
series.  Its  possible  origin  in  the  small  intestine  has 
been  considered. 

The  question  of  the  occurrence  of  tumors  in  ^\dld 
animals  seems  fairly  well  settled  when  twenty-five 
examples  of  malignant  epithelial  neoplasms  can  be  dis- 
covered in  fifty-three  hundred  autopsies.  It  is  interesting 
to  note  the  incidence  of  these  tumors  in  wild-  and  park- 
bred  animals.  Exclusive  of  the  parrakeets  there  are 
twenty-one  cancers,  seventeen  in  known  wild-bred,  two  in 
known  park-bred  specimens,  and  two  with  breeding  un- 
certain. The  average  known  duration  of  captivity  of  the 
wild-bred  animals  is  about  four  years,  while  the  two  park- 
bred  animals  lived  eight  and  eighteen  years.  Thirteen  of 
the  twenty-one  cases  were  males,  eight  females.  Adeno- 
carcinoma was  discovered  twelve  times,   simplex  nine 


Fir,.  49.  — BASAL-CELL  CARCINOMA  OF  TONGUE.  BLACK  BEAR 
(URSUS  AMERICANUS).  NOTE  ULCERATION  WHERE  PIECE  HAS  BEEN 
EXCISED.  AND  ALSO  NODULAR  THICKENING  OF  WHOLE  BASE  OF 
TONGUE. 


Fig.    50. — MICROSCOPICAL  APPKARANCE  OF  TIMOR   IN    Fin.   49. 


NEOPLASMS  477 

times,  medullary  and  squamous  each  twice.  Three  tumors 
of  the  pancreas  and  mammary  gland  were  seen  in  which 
fibrotic  or  scirrhus  areas  were  found,  but  in  no  case  was 
there  detected  that  hard  cicatrizing  cancer  so  conomonly 
found  in  the  human  breast.  All  the  interesting  cases  of 
carcinoma  have  been  recorded  in  the  discussion  of  organs 
from  which  they  took  origin.  The  only  case  of  chorion- 
epithelioma  has  been  reported  in  detail  on  page  308.  The 
two  cases  of  mixed  tumors  are  as  follows :  Mixed  tumor  of 
the  thyroid  and  adenocarcinoma  sarcomatodes  in  the 
liver;  they  have  been  discussed  in  detail  on  pages  334 
and  242  respectively. 

Analysis  of  the  incidence  of  tumors  according  to 
organs  is  disturbed  by  the  large  number  of  cases  in 
Psittaci.  Including  this  order  the  first  place  is  taken  by 
the  kidney,  followed  by  the  liver,  uterus,  muscle,  gastro- 
intestinal tract,  bone  and  cartilage,  thyroid,  adrenal  and 
lung  in  this  order.  Curiously  enough,  if  these  birds  be 
subtracted  the  degree  of  organ  susceptibility  to  new 
growths  is  not  greatly  altered.  The  lead  is  still  held  by 
the  kidney,  the  uterus  occupying  the  second  place  and 
then  in  sequence  the  liver,  gastrointestinal  tract,  muscle, 
thyroid  and  adrenal.  Examination  of  the  figures  for 
mammals  shows  the  uterus  to  lead  in  numbers,  followed 
by  the  liver,  thyroid,  and  mammary  gland.  For  the  birds 
the  kidney  takes  the  undisputed  head  of  the  column  with  a 
total  of  twelve  tumors  (27  per  cent,  of  all  avian  tumors) ; 
the  next  figures  are  shown  by  the  liver,  gastrointestinal 
tract  and  muscle. 

ZOOLOGICAL   AND   PATHOLOGICAL   LIST   OF   TUMORS 

MAMMALIA 

Primates  (2) 

Cercopithecidse — Hamadryas  Baboon  {Papio  hamadryas) 

Papillary  adenoma  of  gastric  mucosa 
Cebidae — Brown  Cebus  {Cebus  fatuellus) 

Hypernephroma  of  right  adrenal 


478  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Lemures  (1) 

Lemuridae — Ring-tailed  Lemur  (Lemur  catta) 

Papillary  adenoma  of  prostate 

Carnivora  (17) 

FelidaB — Clouded  Leopard  {Felis  nebulosa) 
Endothelioma  of  pleura 
Caracal  [Felts  caracal) 

Osteochondroma  of  nose 
Lion  (Felis  leo) 

Malignant  adenoma  of  cervix  uteri 
Metastases  to  lung 
Jaguar  (Felis  onca) 

Fibroadenoma  of  uterus 
Fibroadenoma  of  bile  ducts 
Lymphangioma  of  mesentery 
ViverridsB — Indian  Paradoxure  (Paradoxiirus  niger) 
Adenocarcinoma  of  pancreas 
Malayan  Civet  (Viverra  tangalunga) 
Carcinoma  of  lung 
Canidae — Corsae  Fox  (Canis  corsac) 

Adenoma  of  pancreatic  ducts 
Red  Fox  (Canis  vulpes  pennsylvanicus) 
Cystic  adenoma  of  bile  ducts 
Raccoon  like  Dog  (Canis  procyonoides) 

Adenocarcinoma  sarcomatodes  of  thyroid 
Gray  Fox  (Canis  cinereo-argenteus) 

Papillary  cyst  adenoma  of  bile  ducts 
Prairie  Wolf  (Canis  latrans) 

Sarcoma  of  thyroid  region 
Metastases  to  lungs 
Prairie  Wolf  (Canis  latrans) 

Sarcoma  of  thyroid  region 
Procyonidae — Common  Raccoon  (Procyon  lotor) 

Adenoma  of  pancreas 
UrsidaD — Polar  Bear   (Ursus  maritimus) 

Adenocarcinoma  of  adrenals 
Metastases    to    lungs,    lymph    nodes,    diaphragm 
Black  Bear   (Ursus  americanus) 

Medullary  carcinoma  of  breast 
Metastases  to  lungs 
Black  Bear  (Ursus  americanus) 
Epithelioma  of  tongue 
Phocidae — California  Hair  Seal  (Zalophus  calif  or  nianus) 
Hypernephroma  of  adrenal 


NEOPLASMS  479 

RODENTIA  (12) 

Sciuridae — Beeehy's   Spermophile    {Citellus  grammurus   beecheyi) 
Osteoma  of  sternum 
Gray  Squirrel   {Sciurus  carolinensis  pennsylvanicus) 

Hypernephroma  of  kidney 
Woodchuck  {Arctomys  monax) 
Adenoma  simplex  of  liver 
Muridae — Waltzing  Mouse  [Mus  wagneri  rotans) 

Adenocarcinoma  of  thigh  muscles 
White  footed  Mouse  {Peromyscus  leucopus) 

Carcinoma  simplex  of  mammary  gland 
White  footed  Mouse  {Peromyscus  leucopus) 

Spindle  celled  sarcoma  of  leg 
White  footed  Mouse  (Peromyscus  leucopus) 

Carcinoma  of  mammary  gland 
Larger  Egyptian  Gerbille   (Gerbillus  pyramidum) 
Fibrosarcoma  of  shoulder  region 
Heteromyidae — Kangaroo  Rat  {Perodipus  richardsoni) 

Sarcoma  of  urinary  bladder 
Octodontidae — Coypu    Rat    {Myocastor    coy  pus) 

Sarcoma  of  thyroid 
Hystricidje — ^Canada  Porcupine  {Erethizon  dorsatus) 

Chorionepithelioma  uteri 
DasyproctidaB — Azara's  Agouti   {Dasyprocta  azara) 
Squamous  carcinoma  of  larynx 

Proboscidea  (1) 

Indian  Elephant    (Elephas  indicus) 

Leiomyoma,  uterine  cornua  and  fimbria 

Ungulata  (7) 

Equidas — Chapman's  Zebra   (Equus  burchelli  chapmani) 

Fibroma  peritonei  with  sarcomatous  and  osseous 

change  and  metastases  to  lung 
Bovidae — Isabelline  Gazelle  {Gazella  Isabella) 

Osteofibroma  of  jaw  with  mucoid  degeneration 
Nylghaie  {Boselaphus  tragocamelus) 

Fibroma  uteri 
Dorcas  Goat  {Capra  hircus) 

Lymphosarcoma  of  mediastinum  with  metastases 

to  liver,  kidney  and  lymph  nodes 
CervidaB — Common  Deer   {Manama  virginiana) 

Fibroadenoma  of  bile  ducts 
Camelidae — Alpaca   (Lama  pacos) 

Carcinoma  of  liver  or  bile  ducts  with  extension 

to  intestine 
SuidaB — Wild  Boar  (Sus  scrofa) 

Carcinoma  uteri 


480  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Edentata  (1) 

Dasypodidae — Nine-banded  Armadillo   (Tatu  novemcinctus) 
Fibroma  uteri 

Marsupialia  (7) 

Didelphyidae — Common  Opossum  {Didelphys  virginiana) 
Adenoma  of  kidney 
Common  Opossum  [Didelphys  virginiana) 
Adenocarcinoma  of  mammary  gland 
Dasyuridae — Spotted  tailed  Dasyure    (Dasyiiriis  macidatus) 

Adenocarcinoma  of  intestines  with   metastases  to 
lymphatics,  liver,  spleen,  lungs 
Tasmanian  Devil  {Sarcophilus  ursinus) 
Cystic  epithelioma  of  skin  of  thigh 
Peramelidae — Rabbit  eared  Bandicoot  {Thylacomys  lagotis) 

Carcinoma  of  lung 
Macropodidae — Red  Kangaroo   (Macropus  rufus) 
Malignant  papilloma  of  stomach 
Metastases  to  liver,  spleen,  kidney 
Red  Kangaroo  {Macropus  rufus) 
Carcinoma  of  lung 
Mestastases  to  spleen  and  gastric  wall 

AVES 
Passeres  (7) 

Turdidae — American  Robin  {Planesticus  migratorius) 

Hypernephroma  of  kidney,  metastases  to  intestine 
Crateropodidse — Jungle  Babbler  (Crateropus  canorus) 

Adenoma  of  kidney 
Tanagridae — ^Palm  Tanager  (Tanagra  palmarum) 

Lipoma  of  abdominal  wall 
Fringillidae — Saffron  Finch  {Sycalis  flaveola) 
Adenocarcinoma  of  kidney 
Chestnut  eared  Finch  (Amadina  castanotis) 

Adenocarcinoma  of  kidney  with  metastases  to  lung 
Chestnut  headed  Bunting  (Emberiza  luteola) 
Lipoma  of  scalp 
leteridae — European  Blackbird  {Meriila  merula) 

Hypernephroma  of  kidney  region  with  metastases 
to  liver 

Striges  (1) 

Bubonidae — Great  Horned  Owl   (Bubo  virginianus) 
Papilloma  of  duodenum 

Psittaci  (26) 

Loriidae — Musky  Lorrikeet  (Glossopsittacus  concinnus) 
Carcinoma  of  lung 


NEOPLASMS  481 

CacatuidaB — Roseate  Cockatoo  {Cacatua  roseicapilla) 
Lipoma  of  abdominal  wall 
Roseate  Cockatoo   {Cacatus  roseicapilla) 
Multiple  lipomata  of  abdominal  wall 
Psittacidae — Undulated  Grass  Parrakett  {Melopsittacus  undulatus) 

Glioma  of  brain  with  metastases  to  liver 
Undulated  Grass  Parrakeet  (Melopsittacus  undulatus) 

Hypernephroma  of  adrenal 
Undulated  Grass  Pan-akeet  {Melopsittacus  undulatus) 

Papillary  adenoma  of  kidney 
Undulated  Grass  Pai-rakeet  {Melopsittacus  undulatus) 

Cystic  papillary  adenocarcinoma  of  kidney 
Undulated  Grass  Parrakeet  (Melopsittacus  undulatus) 

Adenocarcinoma  sarcomatodes  of  liver 
Undulated  Grass  Parrakeet  (Melopsittacus  undulatus) 

Papillai-y  cyst  adenoma  of  kidney 
Undulated  Grass  Parrakeet  (Melopsittacus  undulatus) 

Adenoma  of  kidney 
Undulated  Grass  Parrakeet  (Melopsittacus  undulatus) 

Adenoma  of  kidney 
Undulated  Grass  Parrakeet  (Melopsittacus  undulatus) 

Adenocarcinoma  of  oviduct 
Undulated  Grass  Parrakeet  (Melopsittacus  undulatus) 

Carcinoma   simplex   of   liver  with  metastases   to 
liver,  spleen 
Undulated  Grass  Parrakeet  (Melopsittacus  undulatus) 

Carcinoma  simplex  of  liver 
Undulated  Grass  Parrakeet  (Melopsittacus  undulatus) 

Multiple  lipomata 
Undulated  Grass  Parrakeet  (Melopsittacus  undulatus) 

Multiple  lipomata 
Undulated  Grass  Parrakeet  (Melopsittacus  undulatus) 

Sarcoma  of  pectoral  muscle  with  metastases  to  liver 

Undulated  Grass  Parrakeet  (Melopsittacus  undulatus) 

Round  cell  sarcoma  in  region  of  liver,  spleen,  kidney 

Undulated  Grass  Parrakeet  (Melopsittacus  undulatus) 

Carcinoma  simi)lex  of  thyroid 
Blue  fronted  Amazon  (Chrysotis  cestiva) 

Adenocarcinoma   (?)   of  pro  ventricle 
White  fronted  Amazon  (Chrysotis  leucocephala) 

Epithelioma  in  peritoneum  ( ?) 
All  Green  Parrakeet  (Brotogerys  tirica) 

Sarcoma  of  pectoral  muscle 
Red  shouldered  Parrakeet  (Palceornis  eupatrius) 

Sarcoma  of  testes 
Red  shouldered  Parrakeet  (Palceornis  eupatrius) 

Sarcoma  of  testes 


482  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

King  Parrakeet  {Apromictus  cyanopygius) 

Sarcoma  of  ovary 
Crested  Ground  Parrakeet  {Calopsitta  novcehollandi<s) 

Lipoma  of  muscle  of  abdomen  and  chest  walls 

ACCIPITRES   (2) 

Falconidae — Red  shouldered  Buzzard  {Buteo  lineatus) 
Retroperitoneal  sarcoma 
Sparrow  Hawk  {Sparverius  sparverius) 
Lipoma  around  cloaca 

COLUMBJE    (1) 

Columbidffi — ^Scaly  Ground  Dove  (Scardapella  squamosa) 

Sarcoma   (spindle)   of  kidney  with  metastases  to 
tibia 

Galli  (1) 

Phasianidae — Wild  Turkey  {Meleagris  gallopavo) 

Papillary  adenocarcinoma  of  ovai-y 

FULICARI^    (1) 

Rallidae — Moorhen  {Gallinula  chloropus) 
Endothelioma  of  clavicle 

An  SERES  (4) 

Anatidae — Red  headed  Duck  {Fuligula  ferina  americana) 
Papillary  adenoma  of  kidney 
Black  Duck  {Anas  obscura) 

Hypernephroma  of  adrenal 
Lesser  Snow  Goose  {Chen  h.  hyperboreus) 

Fibroma   on   clavicle 
Bean  Goose  {Anser  fabalis) 

Myxosarcoma  of  pectoral  muscle 

Struthiones  (1) 

Rheidae — Common  Rhea  {Rhea  americana) 

Cystic  papillary  adenoma  of  duodenum 


SECTION  XVII 
THE  COMMUNICABLE  DISEASES— PART  I 

Tuberculosis. 

Nearly  all  infectious  diseases  have  either  a  limited 
natural  zoological  distribution  or  are  encountered  chiefly 
in  one  order  or  division.  Thus  typhoid  fever  is  peculiar 
to  man,  hog  cholera  to  swine,  foot-and-mouth  disease  to 
cows.  A  second  group,  including  for  example  anthrax,  ,^ 
variola,  hemorrhagic  septicemia  and  malignant  edema,  is  ' 
somewhat  less  specific,  and  may  occur  in  several  varieties. 
There  is  no  more  widespread,  important  infection, 
zoologically,  economically  and  hygienically,  than  tubercu- 
losis, and  it  would  seem  that  all  kinds  of  vertebrates  are 
subject  to  it.  Its  manifestations  too,  are  sufficiently 
similar  to  support  the  idea  that  there  must  have  been 
originally  a  common  ancestor  of  the  viruses,  which  we  now 
designate  separately  by  a  term  to  indicate  their  immediate 
source  (human,  avian,  etc.),  and  moreover  it  has  been 
shown  that  any  of  the  artificially  separated  varieties  or 
subspecies  may  under  certain  circumstances  infect  all 
zoological  families. 

There  is,  however,  a  varying  resistance  to  the  tubercle 
bacillus,  certain  zoological  groups  standing  out  preemi- 
nently as  more  or  less  susceptible  to  it.  There  is  also  a 
tendency  for  each  animal  group  to  present  features  more 
or  less  peculiar  to  itself,  but  nevertheless  the  character- 
istics, both  gross  and  minute,  of  the  disease  caused  by  the 
Bacillus  tuberculosis  are  sufficiently  similar  to  permit 
close  analogy  and  to  establish  a  diagnosis  when  the  bac- 
teria are  found. 

The  data  collected  at  this  Garden  are  well  suited  to 
elucidate  the  susceptibility  of  wild  animals  under  captive 
conditions  and  to  illustrate  the  nature  of  lesions  in  them. 

483 


484   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Perhaps  they  do  not  offer  a  perfect  cross  section  of 
zoological  distribution  of  tuberculosis  because  of  the  pre- 
dominance in  the  display  of  birds,  of  monkeys  and  of 
ungulates,  but  the  figures  will  be  found  significant  in  cer- 
tain respects.  Such  records  cannot  be  compared  with 
those  obtained  for  domesticated  animals  in  farms  or 
breeding  places,  nor  can  our  material  be  used  to  show 
the  incidence  for  individual  species,  such  as  cows,  dogs 
and  the  like,  in  a  manner  commonly  used  in  veterinary 
literature.  Those  who  are  interested  in  this  phase  of  the 
problem  can  find  in  Lubarsch-Ostertag's  Ergehnisse, 
1917-18,  No.  2,  a  summary  by  Eber  of  all  recent  literature, 
covering  nearly  1,700  references,  which  really  amounts  to 
a  review  of  all  modern  knowledge  of  tuberculosis  in  the 
lower  animals.  The  article  does  not,  however,  attempt 
to  compare  or  contrast  the  incidence  per  order  or  family 
in  wild  varieties  since  little  information  is  available  on 
these  points.  There  are  no  reliable  data  concerning  the 
existence  of  tuberculosis  in  the  wild.  It  is  noted  in  the 
report  of  the  British  Tuberculosis  Commission  that 
monkeys  were  received  with  this  disease ;  Eber  mentions 
that  wild  swine  from  a  private  preser^^e  presented 
undoubted  evidence  of  tuberculosis  and  another  similar 
experience  with  pheasants,  but  these  and  other  citations 
can  give  no  proper  estimate  of  exact  conditions  nor  allow 
a  decision  that  the  infection  exists  at  all  under  natural 
environment  for  in  all  cases  the  association  with  human 
beings  or  domesticated  animals  cannot  be  excluded. 
Tuberculosis  is  usually  described  as  a  disease  of 
civilization  and  its  incidence  surely  varies  directly 
with  crowding. 

The  Table. 

Description  of  Table  23.  The  study  of  our  autopsy 
records  was  begun  by  the  preparation  of  the  accompany- 
ing table,  which  is  based  first  upon  the  zoological  classifi- 
cation per  order  with  subdivisions  for  families  where 


THE  COMMUNICABLE  DISEASES  485 

there  are  sufficient  examples  within  important  orders, 
and  second,  upon  percentage  of  total  cases.  For  the 
analysis  of  the  pathological  types,  headings  of  probable 
origin,  gross  anatomical  lesions  and  visceral  distribution 
were  then  made.  The  first  vertical  column  shows  the  total 
autopsies  per  order,  and  where  families  are  given,  for 
each  of  them.  In  three  instances.  Primates,  Ungulata 
and  GalU,  not  all  families  are  represented,  so  that  the 
total  for  the  order  is  greater  than  that  for  the  subdi- 
\dsions  quoted.  It  is  noteworthy  that  we  have  had  no 
tuberculosis  among  nine  families  of  ungulates;  this  will 
be  discussed  later.  The  second  column  gives  the  total 
cases  of  tuberculosis  for  the  order  and  family,  respec- 
tively, followed  by  a  third  line  of  percentages.  For  the 
analysis  of  the  pathology  in  these  animals  all  the  proto- 
cols were  reviewed.  Forty-eight  of  them  being  found 
unsuitable,  deductions  were  made  according  to  the  natural 
group,  thereby  leaving  the  number  shown  in  the  fifth 
column  for  separation  according  to  origin  and  type. 
Analyses  in  the  succeeding  columns  are  made  upon 
number  of  cases  and  not  upon  percentages,  because  of  the 
confusion  arising  from  small  decimals.  The  actual  rela- 
tionships will  be  pointed  out  in  the  notes.  Percentage  is 
not  so  necessary  because  comparisons  and  contrasts  are 
usually  made  with  classes  or  orders  where  the  figures 
are  comparable. 

It  is  generally  conceded  that  the  principal  and  only 
significant  routes  of  origin  for  tuberculosis  are  via  the 
respiratory  and  alimentary  tracts.  The  criteria  upon 
which  to  decide  the  route  that  has  been  followed  are  by  no 
means  definite  and  may  not  be  for  any  given  case 
unexceptionable.  In  birds  the  alimentary  tract  is  con- 
ceded to  be  the  important  one,  while  in  mammals  an 
aerogenic  route  is  believed  to  be  the  rule.  However,  since 
feeding  experiments  have  sho^vn  that  tubercle  bacilli  can 
gain  the  lungs  by  passing  through  the  intestinal  wall  and 
abdominal  lymphatics  without  leaving  gross  traces,  the 


486   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 


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THE  COMMUNICABLE  DISEASES 


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II 

488   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

decision  that  one  or  the  other  route  has  been  taken  may  be 
erroneous,  and  statistics  therefore  can  often  be  fallacious. 
It  is  usually  the  rule  to  assume  that  the  oldest  or  best 
developed  lesions  occur  where  the  originally  settled 
organisms  exerted  their  maximum  effect.  The  questions 
of  infection-path  and  of  original  lesion  not  having  been 
settled  it  is  obvious  that  decision  as  to  the  route  must  be 
in  the  nature  of  an  estimate.  With  these  limitations  in 
mind  I  have  divided  the  cases  into  probable  respiratory 
and  alimentary  origins  according  to  the  following  criteria. 
Where  the  lesions  were  wholly  respiratory  or  Avithin  the 
lymph  glands  of  the  trachea  and  bronchi  the  decision  was 
not  so  difficult.  Predominance  of  the  pulmonary  disease 
with  recent  lesions  in  other  organs  was  taken  to  indicate 
an  aerogenic  origin.  The  chronic  ulcerative  or  fibrous 
pulmonary  lesions  were  also  ascribed  to  those  beginning 
in  the  lungs.  The  alimentary  tract  was  considered  for 
this  purpose  as  beginning  in  the  tonsillar  area  and  ending 
at  the  anus.  This  is  as  I  understand  the  customary  teach- 
ing. When  the  lymph  nodes  of  the  alimentary  area  were 
advanced  in  the  process,  the  intestinal  method  was  held 
responsible.  It  is  of  course  not  to  be  forgotten  that 
organisms  coughed  up  from  the  lungs  and  swallowed  may 
be  responsible  for  lesions  within  the  alimentaiy  system. 
However,  a  predominance  of  intestinal,  splenic,  hepatic 
and  lymphatic  lesions  caused  me  to  place  the  case  with 
those  originating  from  the  alimentary  tract.  Granting 
the  limitations  of  our  knowledge,  of  the  criteria  and  of  my 
own  judgment,  it  is  noteworthy  that  the  results  of  this 
division  of  the  table  are  not  contradictory  to  the  usual 
teaching,  the  most  conspicuous  being  the  predominance  of 
the  alimentary  infection  of  Aves  and  in  the  order 
Primates,  whereas  the  pulmonary  route  has  the  highest 
figures  for  the  Ungulata. 

The  next  subdivision  of  the  table  concerns  the  gross 
pathological  type.  Beginning  with  the  most  acute  form, 
the   acute   miliary,   progression   is   made   in   terms    of 


THE  COMMUNICABLE  DISEASES  489 

clironicity — then  following  in  order  the  massive  caseous 
form  including  caseous  pneumonia,  the  caseous  miliary  or 
nodular  form  so  well  represented  by  the  monkey,  then  the 
fibroulcerative  type  such  as  one  encounters  in  human 
consumption,  including  also  forms  in  which  fibrosis  pre- 
dominates, and  lastly  the  rather  uncommon  pearl  disease. 
This  classification  has  been  relatively  easy  to  follow  and 
can  be  readily  imagined  by  the  reader.  There  are  of 
course  intermediate  cases  or  transition  forms  and  there 
have  been  instances  partaking  of  more  than  one  character. 
The  groupings  present  only  gross  appearances  and,  with 
few  exceptions,  are  not  to  be  taken  as  direct  indications 
of  type  incidence  in  special  groups. 

Visceral  distribution  is  shown  in  the  last  gross  section 
of  the  table;  single  cases  or  unusual  locations  are  not 
tabulated  but  will  be  separately  discussed.  The  visceral 
distribution  is  made  upon  evident  gross  lesions  or  their 
discovery  in  organs  whose  condition  suggested  the  need  of 
microscopic  study  for  confirmation.  The  figures  in  the 
table  will  be  reviewed  first  upon  the  incidence  as  a  whole 
and  then  between  classes  and  orders.  This  will  be  suc- 
ceeded by  an  analysis  of  the  particulars  for  each  order 
and  then  for  each  of  the  pathological  headings. 

Total  and  Class  iNcroExCE. 

The  autopsies  upon  5,365  animals  have  revealed  the 
existence  of  tuberculosis  in  492,  a  percentage  of  9.1.  This 
means  that  lesions  due  to  the  Bacillus  tuberculosis  were 
present,  but  they  were  not  always  the  cause  of  death,  since 
many  specimens  have  been  executed  and  others  have  had 
sufficient  pathology  to  kill,  aside  from  the  tuberculous 
changes.  Deaths  due  to  the  disease  alone  are  difficult  to 
estimate,  but  seem  to  be  about  325  or  6  per  cent.  These 
figures,  while  they  represent  the  total  incidence,  lose  con- 
siderably in  significance  when  the  factors  are  analyzed. 
Mammalian  incidence  is  14.7  per  cent..  Avian  6.2  per  cent., 
but  the  former  is  based  upon  figures  obtained  from  six  of 


490   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

twelve  orders  numbering  1,860  animals,  whereas  the  latter 
represent  the  cases  in  fourteen  of  twenty  orders  num- 
bering 3,505  birds.  The  percentages  are  considerably 
increased  by  high  figures  for  a  few  orders.  Primates, 
Lemures,  Columbje  for  examples.  There  are  missing 
from  the  list  very  few  orders  of  which  we  have  any 
notable  nimaber  of  autopsies,  Marsupialia  and  Herodiones 
being  the  only  important  ones ;  it  would  seem  that  these 
orders  have  a  high  resistance  to  the  disease. 

Investigation  into  the  origin  of  the  disease  in  mam- 
mals and  birds  shows  wdth  definiteness  the  preponderance 
of  the  alimentary  route  influence  in  the  latter,  but  for  the 
former  the  figures  cannot  be  said  to  be  conclusive.  The 
bird  excretes  large  numbers  of  bacilli  with  the  feces 
thereby  soiling  the  feed  and  the  ground.  This  is  due  to 
the  frequency  of  intestinal  open  lesions  and  to  the  really 
enormous  numbers  of  bacilli  which  are  in  the  morbid  tis- 
sue. I  think  it  can  be  said  with  safety  that,  other  things 
being  equal,  the  bird  excretes  bacilli  constantly  and  in 
greater  numbers  than  does  the  mammal,  and  that 
in  physically  comparable  lesions  there  are  more  bacilli  in 
the  avian  than  in  the  mammalian. 

The  inconclusive  figures  for  the  origin  of  the  disease 
in  the  mammal  can  be  clarified  very  little  by  the  subtrac- 
tion from  the  tables  of  the  figures  for  the  very  susceptible 
Primates.  By  doing  this  it  would  seem  that  the  respira- 
tory route  dominates  as  43  to  29,  whereas  if  the  reasonably 
susceptible  Lemures  be  also  deducted  the  ratio  becomes 
as  38  to  15.  It  would  seem  that  the  evidence  favors  the 
aerogenic  route  in  the  mammal. 

Pathological  Types. 

An  inquiry  into  the  gross  pathological  types  reveals 
at  once  the  frequency  with  which  the  nodular  and  massive 
caseous  forms  appear.  If  the  number  of  cases  be  reduced 
to  percentage  it  will  be  found  that  59  per  cent,  of  all 
specimens  presented  the  nodular  variety  and  26.6  per 


THE  COMMUNICABLE  DISEASES  491 

cent,  the  massive  caseous  form.  These  large  figures 
(equaling  when  combined  85  per  cent.)  coupled  with  the 
fact  that  only  12  per  cent,  of  the  total  were  fibroulcera- 
tive  and  1.8  per  cent,  of  the  pearl  type,  would  seem  to 
indicate  that  the  nodular  and  massive  caseous  processes 
are  the  lesions  to  be  expected  in  wild  animals.  Further- 
more, if  these  nodular  and  caseous  forms  speak  for  recent 
infection  or  acuteness  of  the  morbid  process,  it  would 
seem  that  wild  animals  have  a  low  tissue  resistance 
to  tuberculosis.  It  is  a  widespread  belief,  in  some  degree 
well  supported,  that  a  disease  new  to  an  animal  species  is 
highly  fatal  and  that  the  survival  of  the  race  depends 
upon  an  active  self -immunization  or  the  survival  of  the 
pathologically  least  susceptible.  If  tuberculosis  be  a 
disease  of  civilization,  these  figures  would  suggest  that 
it  is  absent  in  nature.  As  a  further  support  of  this  idea 
it  can  be  said  that  with  the  exception  of  two  cases  in 
ungulates,  no  fibroid  tuberculosis,  approaching  the 
quiescent  type  as  seen  in  man  and  rarely  in  domestic 
animals,  was  encountered.  Very  rarely  calcareous 
deposits  will  be  found  in  both  simian  and  ungulate 
lesions  but  these  need  not  indicate  a  tendency  to  general 
healing  although  at  that  place  the  process  may  be  inac- 
tive. The  bird  uses  considerable  fibrous  tissue  in  the 
construction  of  its  tuberculous  mass  but  fibrosis  never 
masters  the  situation  with  the  formation  of  scar  tissue 
sufficient  to  wall  off  the  process.  Pearl  disease,  a  fibro- 
caseous  condition,  is  not  a  healing  fibroid  procedure  and 
is,  in  our  material,  of  no  numerical  significance. 

Visceral.  Distribution. 

The  distribution  of  the  morbid  lesions  in  the  viscera 
presents  some  interesting  features.  In  the  first  place  the 
data  leave  no  doubt  that  the  most  susceptible  tissue  in 
the  wild  animal  body  is,  as  in  the  case  of  human  and 
domestic  animals,  the  lung.  The  susceptibility  of  this 
organ  in  the  two  classes  is  however  a  different  matter 


492   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

since  in  the  mammal  91.4  per  cent,  show  pulmonary  lesions 
while  only  53.7  per  cent,  of  birds  are  so  affected.  Part  of 
the  reason  for  this  appears  in  the  figures  for  the  principal 
abdominal  organs,  of  which  the  liver  and  spleen  occupy 
the  prominent  places.  The  mammalian  livers  show  58.2 
per  cent.,  the  avian  80.3  per  cent. ;  67.5  per  cent,  of  mam- 
malian spleens,  68.0  per  cent,  of  avian  spleens  have  tuber- 
culous lesions.  The  figure  for  the  mammalian  spleen  is 
distorted  because  of  the  peculiar  susceptibility  of  this 
organ  in  the  monkey,  it  being  conspicuously  free  of 
lesions  in  most  mammalian  orders.  The  intestines  pre- 
sented discernible  lesions  in  practically  40  per  cent,  of 
birds  but  only  in  13.5  per  cent,  of  mammals.  Renal 
involvement  was  found  in  38.4  per  cent,  of  mammals  and 
22.9  per  cent,  of  birds. 

A  study  of  the  changes  in  the  serous  surfaces  is  com- 
plicated by  the  difference  of  anatomy  in  the  two  classes. 
The  mammal  has  separate  closed  serous  sacs  well  guarded 
against  invasion  from  mucous  surfaces  whereas  in  the 
bird  the  air  sacs  and  serous  cavities  are  closely  related, 
the  latter  being  loosely  applied  to  viscera  they  are 
intended  to  cover.  Moreover  in  Aves  direct  infection  of 
the  air  sacs  seems  a  definite  possibility.  Notwithstanding 
the  fact  that  the  bird's  sacs  and  serous  surfaces  appear 
so  open  to  infection  there  is  no  great  preponderance  of 
lesions  within  them — Aves  39.3  per  cent.  Mammalia  32 
per  cent.  The  lymph  nodes  were  tabulated  as  a  tissue 
rather  than  according  to  location,  the  latter  method  be- 
ing found  profitable  for  discussion  in  a  few  orders  only. 
As  might  be  expected  the  abundant  lymph  nodes  of  the 
mammal  were  affected  out  of  all  proportion  to  those  in 
the  bird.  These  matters  will  be  discussed  later.  The 
remaining  figures  on  the  table  have  no  comparative  value. 

Ordinate  Characteristics. 

The  Primates  as  an  order  have  shown  low  resistance 
to  tuberculosis,  a  fact  well  known  to  general  observation. 
How  much  this  is  due  to  the  unsanitary  surroundings  to 


THE  COMMUNICABLE  DISEASES  493 

which  these  naturally  free  active  beasts  are  subjected,  in 
catching,  transporting  and  storing  for  sale,  must  at  pres- 
ent remain  conjectural,  but  they  are  probably  infected 
with  ease  as  our  experience  in  this  laboratory  suggests. 
Desiring  a  tuberculous  monkey  for  certain  tuberculin 
tests,  I  injected  one  hundredth  of  a  milligram  of  a  human 
culture,  known  to  produce  definite  lesions  in  rabbits ;  the 
animal  died  in  three  months  with  advanced  general  tuber- 
culosis. The  unexpected  and  interesting  feature  of  our 
figures  is  the  susceptibility  of  different  families  within 
the  order.  The  Old  World  monkeys,  Simiadae  and  Cerco- 
pithecidse  have  a  combined  incidence  two  and  one-half 
times  as  great  as  the  New  World  Cebida,  and  the  marmo- 
sets had  no  tuberculosis  at  all  in  the  thirty-two  speci- 
mens. Possibly  this  is  a  matter  of  transportation  and 
handling,  which  reduces  the  resistance  and  offers  chance 
to  infect  apes  and  baboons.  The  New  World  capucin  mon- 
keys have  their  exposure  too  since  many  of  them  are 
household  pets  before  the  Garden  receives  them.  Their 
usual  life  in  captivity  is  however  shorter  than  that  of 
Old  World  varieties,  they  therefore  being  exposed  to  in- 
fection for  a  shorter  time.  It  would  seem  however  that 
American  Primates  are  more  resistant  to  the  disease  than 
African  and  Australasian. 

The  form  of  tuberculosis  to  which  this  order  is  liable 
is  well  described  in  text-books,  it  being  so  characteristic 
that  the  term  ''  monkey  tuberculosis  "  is  used  to  dis- 
tinguish it.  The  purpose  of  the  term  is  to  compare  the 
lesions  with  certain  cases  of  generalized  tuberculosis  in 
children.  It  is  characterized  by  a  nodular  involvement  of 
the  liver  and  of  the  spleen  particularly,  sometimes  also 
of  the  lungs  but  in  fatal  cases  the  last  organ  is  commonly 
the  seat  of  massive  caseation  or  caseous  pneumonia.  The 
prominence  of  the  pulmonary  lesions  often  makes  a  deci- 
sion of  origin  difficult  since  important  changes  may  be 
found  in  the  liver  and  abdominal  nodes.  Blair  at  New 
York,  and  Rabinomtsch  at  Berlin  are  of  the  impression 
that  many  cases  of  monkey  tuberculosis  start  by  pharyn- 


494  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

geal  and  tonsillar  infection  because  they  found  cervical 
adenitis  so  commonly.  Our  records  and  specimens  would 
support  this  idea  in  only  fourteen  instances  and  I  am  of 
the  impression  that  the  lower  intestinal  route  is  more 
often  responsible,  even  to  a  higher  figure  than  is  recorded 
in  the  table.  This  view  is  based  upon  the  frequent  occur- 
rence of  enlarged  glands  in  the  mesentery,  retroperito- 
neum  and  posterior  mediastinum,  in  the  latter  location 
being  quite  as  prominent  if  not  more  so  than  in  the 
bronchial  and  tracheal  area.  Occasionally  deposits  of 
calcareous  matter  will  be  found  in  old  caseous  glands 
but  in  such  animals  there  has  always  been  some  other 
spot  of  activity  of  tuberculosis.  The  frequency  with 
which  the  liver  and  spleen  are  affected  gives  opportunity 
for  hematogenic  spread,  a  method  of  no  small  importance 
in  the  opinion  of  Eber.  The  chronic  ulcerative  form  is 
quite  well  displayed  in  monkeys,  interestingly  enough  to 
cite  an  illustrative  case  at  the  end  of  this  division  of  the 
discussion.  Five  cases  of  distinct  chronic  cavitation 
were  encountered;  several  small  recent  cavities  were 
found  in  the  massive  caseous  pneumonic  cases.  The  two 
acute  miliary  cases  and  one  of  the  pearl  type  will  be  dis- 
cussed briefly  on  a  later  page. 

Despite  the  prominence  of  the  liver,  spleen  and  lymph 
nodes,  the  lungs  stand  ahead  of  all  others  by  a  safe  margin 
of  visceral  incidence.  The  spleen  stands  in  the  third 
place  in  this  order  and  in  the  next,  Lemures,  but  in  no 
other  manomalian  group  does  this  organ  occupy  so  promi- 
nent a  position.  The  susceptibility  of  l}Tnphatic,  tissue  in 
the  monkey  is  further  illustrated  by  the  large  number  of 
of  cases  showing  lesions  in  lymph  nodes.  It  is  rather 
striking  however  that  our  material  showed  very  few 
active  ulcerations  in  the  hTQphoid  plaques  of  the  intes- 
tinal wall  nor  indeed  do  the  intestines  present  a  large 
numerical  involvement.  Serous  membrane  tuberculosis  is 
chiefly  that  of  the  pleura,  upon  which  early  precaseous 
tubercles  are  frequently  found,  usually  in  conjunction 


THE  COMMUNICABLE  DISEASES  495 

with  pulmonary  disease.  Tuberculous  peritonitis  of  the 
plastic  and  nodular  variety  occurred  only  six  times, 
though  light  adhesions  to  nodules  in  the  liver  and  spleen 
were  quite  common.  Pericarditis  was  found  five  times, 
in  three  of  which  there  was  myocardial  disease ;  which  of 
the  two  was  primary  was  not  indicated  in  the  notes  but 
from  present  reading  it  would  seem  that  the  heart  muscle 
was  involved  first. 

The  next  order,  Lemures,  has  a  susceptibility  of  about 
half  that  of  the  monkeys  judging  by  the  percentage  inci- 
dence. Analytically  the  members  of  this  group  react  quite 
like  the  preceding  order  in  having  the  same  types  of 
origin,  pathology  and  organic  distribution.  Since  they 
are  so  close  zoologically,  present  similar  lesions  and  are 
tested  in  the  same  manner  as  the  monkeys,  we  group  them 
together  and  shall  proceed  to  discuss  special  cases  of 
interest  in  both  orders. 

Special  Cases  in  Primates  and  Lemures.  Acute  gen- 
eral miliary  tuberculosis  occurred  thrice  in  Old  World 
and  once  in  New  World  monkeys.  The  first  case  took  its 
origin  in  a  caseous  gland  in  the  bronchotracheal  area, 
the  lung  showing  a  minor  degree  of  involvement  with 
milia  but  no  older  process.  The  second  took  its  origin  in 
the  mesenteric  area  and  the  organs  of  this  section  of  the 
body  were  most  affected.  The  only  case  in  the  Cebidas 
seemed  to  be  of  intestinal  origin  since  an  acute  plastic 
peritonitis  with  fluid  exudate  accompanied  the  generally 
miliary  disease. 

Pearl  disease  of  the  bovine  type  has  been  encountered 
on  four  occasions  but  it  cannot  be  said  to  have  developed 
to  the  state  of  perfection  seen  in  the  cow.  There  is  lack- 
ing the  masses  of  nodules  growing  together  in  a  fungoid 
character  usually  seen  on  the  pleura  and  peritoneum.  The 
monkey  form  is  in  isolated  nodules  of  gray  yellow  color 
which  may  show  caseous  centres.  The  visceral  lesions 
are  in  firm  separate  areas  not  tending  to  soften  or  coa- 
lesce. Two  of  these  monkeys  died  from  recent  pulmonary 


496   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

exacerbations.  From  one  of  the  cases  a  bovine  bacillus, 
judging  from  culture  and  rabbit  virulence,  was  isolated. 
As  a  good  example  of  monkey  tuberculosis,  illustrating 
at  the  same  time  a  chronic  ulcerative  pneumonitis  with 
cavitation,  the  following  case  is  cited : 

Green  Moakey  {Cercopithecus  callitrichus)  9  •  Was  coughing  and 
drooping  for  two  days  before  death.  Chronic  ulcerative  tuberculosis 
of  lungs  with  cavity  formation;  early  conglomerate  tuberculosis  of  liver; 
conglomerate  caseous  tuberculosis  of  spleen;  early  conglomerate  tuber- 
culosis of  ileum  (Peyer's  patches);  acute  catarrhal  enteritis;  miliary 
tuberculosis  of  right  kidney.  The  animal  is  well  preserved,  sleek,  with 
a  moderate  amount  of  fat.  The  left  pleura  is  largely  obliterated  by 
adhesions  in  the  lower  portion.  As  lung  is  freed  it  is  torn,  showing  a 
cavity  measuring  3x3x4  cm.,  which  is  filled  with  a  curdy  gray  material. 
Cavity  has  well  defined  walls.  Rest  of  lower  lobe  in  which  this  cavity  lies 
is  solidified,  red  and  edematous  and  contains  numerous  conglomerate 
tubercles.  Upper  lobe  practically  free  of  tubercles;  shows  compensa- 
tory emphysema.  The  right  lung  closely  resembles  the  left  but  lacks 
the  cavity.  The  liver  is  enlarged,  soft  and  friable,  of  red  color,  spotted 
yellow.  Serous  and  section  surfaces  show  closely  packed  early  con- 
glomerate tubercles.  The  spleen  is  of  normal  size,  soft,  has  red  pulp 
with  large  conglomerate  tubercles  which  project  slightly  on  the  capsule. 
The  kidneys  are  apparently  normal  except  for  the  presence  of  two  or 
three  subcapsular  large,  solitary  tubercles  in  the  right  organ.  The 
duodenum  has  thickened  walls,  mucosa  bright,  brilliant  scarlet  hue.  In 
the  ileum  the  walls  are  thickened,  mucosa  bright  red,  agminated  follicles 
hyperplastic  elevated  and  display  several  (4-12)  miliary  tubercles.  These 
may  be  seen  shining  through  on  the  serous  surface  but  there  is  no 
peritoneal  tuberculosis.  No  ulceration  of  Peyer's  patches.  Contents 
of  large  intestine  is  rather  dry  and  here  the  mucosa  shows  exaggerated 
rugae  which  cannot  be  smoothed  out.  Walls  are  thickened,  and  ulcera- 
tion, while  suggested,  cannot  surely  be  determined. 

An  interesting  case  of  primary  tuberculosis  in  the 
laryn:x  detected  at  postmortem  after  a  tuberculin  injec- 
tion is  as  follows : 

Black  and  White  Lemur  {Lemur  varius)  6  .  Miliary  tuberculosis 
of  larynx;  perilaryngitis  and  retropharyngeal  lymphadenitis.  Killed 
because  of  unsatisfactory  chart  after  injection  of  tuberculin.  The  only 
tuberculous  lesion  to  be  found  in  the  body,  which  is  in  excellent  shape, 
is  in  and  about  the  larynx.  The  lesions  within  are  on  the  epiglottis, 
false  and  true  vocal  cords  and  the  main  ventricle.  On  either  side  of  the 
root  of  the  epiglottis,  there  are  a  few  recent  tubercles.  The  lesion 
in   the   retropharyngeal   lymphatics   is   recent   and   diffuse.     This   is 


Fig.  51. — BOVINE  TUBERCULOSIS  IN  THE  MONKEV.  THIS  SPECIMEN  >H()\VS  THE  BOVINE 
PEARL  DISE.iVSE  ON  THE  COSTAL  PLEURA,  AND  SERVES  AS  WELL  TO  ILLlS'IR-VIE  THE  NODULAR 
TUBERCLES  COMMONLY   FOUND  IN  THE  SPLEEN   OF  ALL  PRIMATE    TUBERCULOSIS. 


THE  COMMUNICABLE  DISEASES  497 

probably  primary  as  it  is  not  known  that  the  retropharyngeal  glands 
drain  to  or  from  the  larynx.  About  the  lesions  on  the  laryngeal  mucosa 
there  is  an  area  of  congestion  probably  due  to  the  tuberculin  injection. 

Local  lymphatic  tuberculosis  of  comparative  interest 
was  encountered  a  few  times.  Three  cases  of  cervical 
adenitis,  large  enough  to  be  visible,  were  seen,  of  which 
one  broke  down  about  two  weeks  before  death,  and  dis- 
charged. The  others  did  not  ulcerate  through  the  skin 
but,  contrary  to  the  usual  rule  for  the  human  being, 
remained  as  isolated  glands  only  lightly  adherent  to  one 
another  where  they  lay  adjacent.  There  was  also  seen 
an  ulcerating  tuberculous  lymph  node  in  the  groin  of  one 
monkey,  the  animal  ha\dng  rather  pronounced  abdominal 
and  pelvic  tuberculosis.  Two  instances  of  tonsillar  tuber- 
culosis are  recorded,  in  both  of  which  the  lesion  was  of 
some  duration  and  associated  with  caseation  in  the  lymph 
node  lying  immediately  behind  and  below  it.  A  Guinea 
Baboon  {Papio  sphinx)  had  as  an  unusual  part  of  his  gen- 
eral tuberculosis,  an  active  caseopurulent  collection  in  the 
antrum  of  Highmore,  which  attacked  the  upper  maxilla 
and  immediately  adjacent  muscle ;  tubercle  bacilli  could  be 
demonstrated. 

One  of  the  cases  of  nodular  or  massive  peritoneal 
tuberculosis  is  quite  like  the  tumor-forming  variety  of 
human  adolescents ;  it  is  as  follows : 

Reddish  Macaque  (Macacus  rufescens).  Caseous  tuberculosis  of 
mesentery  and  spleen;  miliary  tuberculosis  of  lungs,  pleura  and  liver; 
chronic  myocarditis.  The  lymphatic  glands  of  the  posterior  and  supe- 
rior mediastinum  and  bronchi  are  slightly  enlarged,  soft  and  anthracotic 
but  do  not  show  any  tuberculous  change.  Both  lungs  are  riddled  with 
small,  firm,  gray  miliary  tubercles,  some  surrounded  by  a  clear  mantle 
of  connective  tissue.  The  intervening  lung  is  practically  normal.  Pleura 
over  base  of  right  lung  on  both  surfaces  shows  small,  pale  miliary 
tubercles.  The  liver  contains  various  sized  miliary  tubercles.  There 
is  a  large  caseous  mass  in  posterior  end  of  spleen  with  adhesions  to 
kidney,  stomach  and  colon.  Retroperitoneal  glands  are  much  enlarged, 
firm,  homogeneous — probably  tuberculosis  of  a  different  type.  Lesser 
omentum  contains  one  caseous  gland.  Few  caseous  glands  in  great 
omentum.  In  the  right  iliac  region  there  is  a  large  mass  involving  many 
coils  of  intestine.     It  is  found  to  arise  probably  from  the  ileocecal 


498  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

glands  and  can  be  traced  along  the  mesentery  to  the  central  lymphatic 
stalk.  The  mass  involved  the  tissues  of  the  mesentery  and  surrounds 
many  coils  of  intestine.  Cecum  and  first  part  of  colon  can  be  traced 
over  its  right  side.  Rectum  is  free  except  on  right  side  where  it  is 
lightly  attached  to  the  mass.  Epicardium  is  grray  and  irregularly 
thickened  and  the  muscle  just  beneath  serous  membrane  is  pale  and 
streaked  with  red  lines. 

There  have  been  three  cases  of  tuberculosis  of  the 
internal  male  genital  area,  one  of  which  was  suspected  of 
having  been  the  primary  seat  of  the  disease ;  it  was  de- 
scribed on  page  315.  The  other  two  could  have  been  sec- 
ondary since  other  points  of  morbid  change  were  as  old 
or  older.  One  of  these  cases  formed  a  tumor  as  large  as  a 
goose  egg  at  the  vesical  neck,  obstructing  the  flow  of  urine 
and  blocking  up  the  seminal  vesical,  in  consequence  of 
which  paralytic  distention  occurred  in  the  bladder  while 
the  vesicles  were  tightly  filled  with  inspissated  semen.  A 
case  of  Fallopian  salpingitis  has  also  been  cited.  Two 
instances  of  cerebral  tuberculomata  have  already  been 
described. 

Camivora.  This  order  has  the  reputation  of  being 
quite  resistant  to  the  tubercle  bacillus,  based  upon  the 
relative  infrequency  among  cats  and  dogs  in  contrast  to 
cows  and  swine.  Some  veterinary  statistics  cite  the 
incidence  up  to  5  per  cent.,  and  occasional  references 
may  be  found  to  tuberculosis  in  circus  lions  and  tigers. 
Our  records  would  suggest  that  in  gardens  the  wild 
varieties  of  this  order  have  about  as  much  of  the  infection 
as  the  domestic  carnivores,  3.5  per  cent.  The  group  is 
made  of  six  FelidaB,  one  Viverridae,  three  Canidae,  six 
Procyonidae,  and  one  Ursidae.  The  first  family  includes  a 
lion,  tiger,  a  jaguar  and  three  smaller  cats.  The  Canidae 
are  all  small  foxes.  The  Procyonidas  are  all  coatis.  The 
features  of  this  order  are  the  occurrence  of  the  fibro- 
ulcerative  variety  with  cavitation  in  the  Felidae  and  the 
caseous  nodular  abdominal  and  glandular  disease  in  the 
coatis.  All  these  animals,  even  those  of  the  last  named 
variety  and  pathological  type,  tend  to  show  some  tissue 


THE  COMMUNICABLE  DISEASES  499 

resistance  to  the  tuberculous  disease.  Connective  tissue 
activity  is  characteristic  of  the  process,  considerable  dis- 
tortion being  produced  by  the  fibrosis.  This  feature  is 
borne  out  where  the  tissues  are  studied  microscopically. 
Definite  milia  are  sometimes  found,  but  they  consist 
of  epithelioid  and  round  cells  with  imperfect  caseation, 
giant  cells  being  often  missing.  About  the  miliary 
tubercles  a  diffuse  and  not  essentially  specific  tuberculous 
granulation  tissue  is  found,  mixed  with  which  is  much 
connective  tissue  growth.  The  fibrotic  adhesion-form- 
ing serous  membrane  tuberculosis  of  carnivores  seems 
worthy  of  emphasis  by  the  citing  of  a  case  in  point. 
White-nosed  coati  {Nasua  narica),  was  received  in  poor 
condition  and  died  in  a  few  days.  Upon  dissection  a 
slightly  turbid  yellowish  fluid  was  found  to  occupy  what 
remained  of  the  peritoneal  cavity  which  was  reduced  in 
size  by  dense  adhesions  of  the  intestines  into  an  inflam- 
matory mass.  The  omentum  was  a  diffuse  thickened 
apron,  also  beset  with  fine  tubercles,  lying  over  the  mass. 
Fine  young  tubercles  could  also  be  found  upon  the 
intestines  and  liver  while  the  mesenteric  lymph  nodes 
were  early  in  caseation ;  thoracic  organs  not  infected. 

One  of  the  most  interesting  cases  concerned  hyper- 
trophic osteitis  in  a  chronically  tuberculous  lion ;  the  feet 
are  discussed  on  page  346.  This  process  was  described 
by  Marie  for  human  beings  many  years  ago,  and  was 
reported  in  dogs  by  Cadiot(l)  in  1912.  This  beast  was 
one  of  five  large  cats  which  have  died  from  tuberculosis 
in  its  chronic  ulcerative  form.  Three  of  the  cats,  one  fox 
and  the  bear  showed  definite  cavitations  of  a  ragged  locu- 
lated  form.  The  cavities  were  usually  of  the  multiple 
variety  and  were  found  in  the  posterior,  that  is  lower  lobe. 

Rodentia.  The  paucity  of  cases  in  this  order  permits 
little  information  to  be  drawn  from  the  form  of  tubercu- 
losis. The  total  seems  to  have  been  swollen  by  a  group  of 
three  beavers,  all  of  which  came  in  one  shipment.     The 

(1)   Rev.  de  Med.  Vet.  T.    89,  p.  221. 


500   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

remainder  were  a  Capybara  and  an  Agouti.  The  general 
type  is  that  of  much  caseation  with  little  or  no  sur- 
rounding fibrosis.    One  illustrative  case  is  cited : 

American  Beaver  {Castor  canadensis).  General  tuberculosis.  The 
animal  presents  generalized  tuberculosis.  The  regional  lymph  nodes 
show  caseous  nodules.  The  right  hip  joint  shows  caseous  material 
about  the  acetabulum  with  necrosis  and  pathological  fractures  in 
the  OS  innominatum  immediately  above  the  acetabulum  and  including 
its  cavity.  Lungs  show  almost  no  normal  respiratory  tissue,  the  proc- 
ess being  a  diffuse  precaseous,  partly  gelatinous  pneumonic  phthisis. 
The  superior  and  posterior  mediastina  show  caseous  glands.  There  is 
miliary  tuberculosis  of  the  liver.  Nodular  caseous  tuberculosis  of  the 
spleen  with  small  tubercles  and  some  cirrhosis  of  the  intervening  tissue. 
There  are  caseous  nodules  in  all  perirenal  glands  and  in  the  kidney 
cortices.  The  psoas  muscle  glands  are  densely  caseous.  The  pelvic 
organs  except  about  the  right  acetabulum  escape  involvement.  Adre- 
nals not  opened  but  probably  not  involved. 

Ungulata.  This  order  shows  the  most  definite  figures 
among  those  for  the  order  of  mammals.  Nearly  one- 
tenth  of  the  whole  number  of  specimens  have  had  some 
form  of  tuberculosis  and  of  a  very  definite  character.  It 
is  well  at  first  to  mention,  however,  that  only  four  of 
thirteen  families  are  represented,  from  which  four  came 
328  of  the  total  365  autopsies.  The  remaining  thirty- 
seven  were  such  animals  as  tapirs,  giraffes,  swine,  and 
peccaries,  in  all  of  which  tuberculosis  has  been  reported 
from  elsewhere. 

Pulmonary  disease  with  less  prominent  lesions  in 
other  organs,  especially  the  intestines  and  their  related 
glands,  speaks  in  favor  of  the  aerogenic  route  being  the 
common  one.  This  of  course  has  been  a  bone  of  conten- 
tion among  veterinarians,  and  I  do  not  presume  to  settle 
the  matter  with  these  figures. 

This  order  resists  tuberculosis  to  a  certain  degree  as 
attested  by  the  fibroulcerative  character  of  the  majority 
of  the  cases.  Two  instances,  one  in  a  buffalo  and  one  in 
a  deer,  showed  very  highly  fibrotic  puhnonary  lesions 
with  a  partial  attempt  to  surround  and  wall  off  numerous 
areas  of  caseation.    So  too  in  this  order  there  is  a  greater 


THE  COMMUNICABLE  DISEASES  501 

tendency  to  calcification,  both  in  the  intra-  and  extra- 
pulmonary nodes.  It  is  to  be  emphasized  that  in  our 
material  the  thoracic  lymph  nodes  are  affected  more 
than  the  abdominal  and  regional  as  3  to  1.  The  apparent 
immunity  of  the  spleen  of  this  order  is  well  illustrated. 

The  paucity  of  serous  surface  involvements  in  the 
wild  BovidaB  and  their  prominence  in  the  Cervidae  cannot 
be  ignored  in  the  figures,  but  it  seems  misleading  since 
pleural  growths  and  adhesions  are  quite  common  in  the 
domestic  Bovidae.  The  case  in  the  Equidae  was  that  of  a 
Zebra  with  a  large  tuberculous  abscess  in  the  retro- 
peritoneal glands  forming  a  tumor  in  the  left  renal  region. 
It  was  quite  well  surrounded  by  fibrosis,  and  the  infection 
had  not  extended ;  it  seemed  quite  recent.  Analysis  of  the 
figures  for  the  remaining  three  families  of  ungulates 
offers  little  for  contrast  and  much  for  comparison;  it  is 
the  usual  picture  as  seen  in  the  domestic  cow.  Some 
special  cases  are  worthy  of  review. 

An  interesting  specimen  of  softened  glands  chiefly  on 
one  side  of  the  neck  was  found  in  a  Fallow  deer  {Cervus 
dama).  It  resembled  the  juvenile  human  cases  that 
require  surgical  attention.  Although  palpable  lymph 
nodes  can  be  found  in  practically  all  cases  of  generalized 
tuberculosis  in  the  Ungulata,  this  is  the  only  case  in  our 
records  in  which  they  have  presented  a  large  tumefaction 
and  broken  down.  Pulmonary  cavitation  is  recorded  but 
thrice,  one  for  each  of  the  last  three  families.  Fibro- 
caseous  tuberculosis  of  the  testes  was  discovered  in  a 
Nylghaie  {Boselaphus  tragocamelus),  but  there  is  no 
knowledge  of  mating  or  offspring.  An  ischiorectal 
abscess  was  found  in  an  American  Bison  {Bison  bison) 
showing  nodular  precaseous  tubercles  of  the  lung.  The 
former  was  the  cause  of  death.  No  tubercle  bacilli  could 
be  found  in  the  abscess  contents,  so  that  the  tuberculous 
basis  is  inferred,  not  proven.  Tuberculous  salpingitis 
in  a  Nylghaie  was  discussed  on  page  306. 


502  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Proboscidea.  Eber  mentions  in  the  article  already 
referred  to  that  there  are  three  reports  in  the  literature  of 
tuberculosis  in  elephants.  When  looking  for  an  explana- 
tion of  tuberculosis  in  this  animal  it  must  be  remembered 
that  it  is  one  of  the  most  attractive  objects  in  a  zoological 
garden  and  receives  perhaps  more  attention,  including 
feeding,  from  visitors  than  any  other  specimen.  The  beast 
while  possessing  some  tissue  resistance  to  tuberculosis, 
is  by  no  means  immune  thereto,  as  has  been  thought  by 
some  persons  on  account  of  its  reputed  longevity,  and 
therefore  he  is  to  be  protected  from  infection  just  as 
much  as  other  animals.  It  would  appear  that  he  may 
present  caseous  pneumonia  or  nodular  caseous  dissemi- 
nated lesions.  Our  two  cases,  in  animals  at  the  Garden 
twenty  and  thirty-eight  years  respectively,  were  both  of 
the  fibrocaseous  variety;  the  lesion  was  confined  to  the 
lungs.    A  brief  description  of  their  lesions  is  as  follows : 

Indian  Elephant  {Elephas  indicus)  6  .  Chronic  polyarthritis. 
Chronic  myocarditis.  Chronic  hepatitis  (cirrhosis).  Parenchymatous 
nephritis.  Chronic  tuberculosis  of  the  lungs,  partly  encapsulated.  Pig- 
mentation of  the  spleen.  The  pleurae  are  very  fat  but  the  surfaces  are 
smooth  and  devoid  of  adhesions.  The  lymph  nodes  of  the  mediastinum 
are  about  10  x  20  cm.  for  the  largest  while  the  smaller  ones  vary  around 
2x4  cm.  They  are  firm,  deep  red-brown  Avithout  clear  divisions  into 
medulla  and  follicular  cortex.  There  are  several  large,  firm,  pale  rather 
cheesy  follicles  in  all  the  large  ones  and  a  few  of  the  small.  These  do 
not  appear  like  tuberculosis.  The  lungs  are  flaccid  and  soft;  gray 
and  red  mottled.  The  bronchi  are  firm  and  stand  open.  Around  one 
in  the  upper  lobe  of  the  right  lung,  there  is  a  large  area  of  cheesy 
degeneration  around  which  a  zone  of  connective  tissue  has  formed. 
This  extends  about  the  bronchus  about  halfway  in  a  sheath-like  man- 
ner. There  is  also  a  separate  nodule  the  size  of  a  cherry  with  a  cheesy 
centre.  The  trachea  appears  normal.  Tubercle  bacilli  could  be  demon- 
strated in  the  cheesy  material.  Microscopic  section  of  lung  around  the 
cheesy  area  shows  a  low-grade  chronic  granulation  tissue  in  some  places 
enclosing  cheesy  masses  with  giant  cells  on  the  margin.  The  neighbor- 
ing septa  are  slightly  thickened  and  in  some  places  broken,  fonning 
emphysematous  cavities.     Some  of  these  cavities  are  edematous. 

Indian  Elephant  {Elephas  indicus)  9.  Miliary  and  conglomerate 
caseous  tuberculosis  of  lung.  Edema  of  lungs.  Endarteritis  deformans 
of  lung.    Cloudy  swelling  of  liver.    Chronic  passive  congestion  of  liver. 


THE  COMMUNICABLE  DISEASES  503 

Hemosiderin  pigmentation  of  liver.  Acute  parenchymatous  nephritis. 
Chronic  passive  congestion  of  spleen.  Chronic  hyaline  perisplenitis. 
Multiple  calcified  fibroid  tumors  of  uterus.  Leiomyoma  of  uterine  cornu. 
Senile  atrophy  of  ovaries.  Acute  catarrhal  enteritis.  There  are  some 
adhesions  of  the  upper  lobes  of  the  lung  to  the  ribs.  The  lungs  are 
large,  increased  in  weight,  color  pink  and  mottled  red,  air  content 
diminished.  There  are  several  masses  of  tubercles,  each  as  large  as 
a  cocoanut,  in  both  lobes.  In  one  such  the  tubercles  are  yellow  and 
caseous ;  some  are  fibroid  but  none  are  liquefied  or  calcified.  The  fibrous 
tissue  of  the  lung  parenchyma  here  is  much  overgrown.  In  one  instance 
the  terminus  of  a  bronchus  is  solidly  plugged  by  caseous  material. 
Mucosa  of  bronchi  is  reddened,  markedly  ulcerated,  ulcers  overlaid 
by  mucopus. 

Individual  Features  of  Avian  Tuberculosis. 

The  avian  form  of  tuberculosis  is  somewhat  peculiar 
iu  its  physical  appearance  as  well  as  in  its  distribution.) 
The  isolated  nodular  type  is  far  and  away  more  common 
by  more  than  100  per  cent,  than  all  the  other  types  com- 
bined. These  nodules  are  usually  well  circumscribed, 
and  to  the  naked  eye  suggest  that  they  have  a  restraining 
fibroid  wall.  This  is,  however,  not  the  case,  the  impres- 
sion being  due  to  the  dense  but  actively  growing  fibro- 
cellular  cortical  zone  of  the  tubercle.  The  centre  of  the 
nodule,  instead  of  having  the  soft  character  like  Camem- 
bert  cheese,  resembles  the  firm  but  brittle  American  dairy 
cheese.  Upon  opening  such  an  area  the  central  necrotic 
mass  may  split  away  from  its  cortex  and  even  shell  out, 
lea\ang  a  cavity  lined  by  a  graj^-yellow  membrane.  These 
characters  are  best  displayed  in  nodules  of  moderate  size, 
the  small  ones  being  like  the  yellow  mammalian  analogue, 
the  large  being  like  indefinite  cheesy  masses.  In  the  sur- 
rounding tissue  evidences  of  inflammatory  processes 
seem  decidedly  greater  in  our  material  than  I  am  accus- 
tomed to  see  in  human  and  veterinary  pathology.  This, 
it  seems,  should  be  emphasized  since  secondary  infection 
with  pus  cocci  and  other  pathogenic  germs  appears  less 
often  in  birds  than  in  mammals. 

The    difference    speaks,   therefore    for    a   difference 
either  in  the  tubercle  bacillus  of  birds  or  the  avian  physi- 


504   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

ology.  Judging  by  the  limited  morbid  processes  produced 
by  injecting  avian  bacilli  into  rabbits  and  guinea-pigs 
the  reaction  of  the  bird  itself  would  not  seem  wholly 
responsible  for  the  difference.  The  local  tissue  reaction 
in  all  avian  lesions  is  mononuclear  and  fibrous,  softening 
and  pus  being  rare.  It  would  seem  from  this  and  similar 
operations  that  the  bird  expresses  its  resistance  to  the 
bacteria  by  a  fibrocellular  reaction  which  goes  on  to 
fibrosis  without  softening;  perhaps  this  means  also  that 
their  polynuclears  are  not  sufficiently  active,  but  the 
pathogenic  power  of  the  bacillus  itself  doubtless 
is  individualistic. 

The  character  of  the  cheesy  degeneration  is  likemse 
different  from  the  mammalian.  It  seems  like  an  abrupt 
hyaline  necrosis  of  a  large  central  mass  and  not  the  slower 
cell  death  seen  in  the  other  types  of  tubercle.  At  times 
the  degenerated  area,  instead  of  having  the  yellowish 
color  of  caseation,  will  present  what  we  have  designated 
''gelatinous  tuberculosis,"  the  whole  infiltrated  area 
resembling  boiled  sago  or  tapioca.  This  seems  to  be  a 
complete  homogeneous  coagulation  or  hyaline  necrosis 
of  the  whole  mass  out  to  the  delicate  fibrous  mantle  sup- 
plied by  the  tissue  in  which  the  tubercle  lies. 

The  organic  distribution  of  tuberculous  lesions  has 
already  received  some  attention  and  is  to  be  discussed 
with  the  orders.  There  are,  however,  some  localities 
affected  conspicuously  in  the  bird.  The  skin  lesions  often 
attract  attention  during  life.  They  occur  around  the  eye,  at 
wing  joints,  on  the  cresta  sterni  and  on  the  legs.  Parrots 
and  jays  have  shown  nodular  or  diffuse  growths  around 
the  eye,  originating  both  in  the  lids  and  orbit,  which  on 
section  have  proved  to  be  tuberculous.  These  seldom 
ulcerate,  but  those  upon  the  skin  of  the  breast  and  wings 
tend  to  have  superficial  erosions  or  deep  ulcers.  The  latter 
lesions  are  more  common  upon  pigeons  but  have  been 
seen  in  Psittaci  and  Galli.     Toucans  and  pigeons  when 


-MASSIVE  TUBERCULOSIS  IN   LIVER   AND  SEVERAL  MURAL  TUBERCLES  OF  INTESTINE 
COMMON   PEA   FOWL   (PAVO   CRISTATA). 


Fig.    53.— nodular   HEPATIC   LESIONS   IN  THE  LIVER   OF  A   DOVK. 


THE  COMMUNICABLE  DISEASES  505 

pinioned,  have  on  three  occasions  shown  a  tuberculous 
mass  on  the  stump. 

Two  parrots  with  hyperkeratosis  of  the  beak  and  of 
the  skin  of  the  feet,  have  also  had  tuberculosis.  These 
have  been  mentioned  in  literature  as  of  tuberculous 
origin.  One  case  well  studied  failed  to  show  tubercle 
bacilli  in  the  corns.  In  the  absence  of  tubercle  bacilli,  one 
is  inclined  to  think  that  this  might  be  explained  on  the 
basis  of  a  circulating  toxin  such  as  is  assumed  to  be 
responsible  for  hypertrophic  periosteitis.  This  latter 
condition  has  not  been  seen  in  birds. 

Still  another  type  of  occasional  occurrence  deserves 
mention.  While  most  of  the  lesions  in  birds  correspond 
to  the  description  given  in  the  preceding  pages, 
some  lesions  fail  to  degenerate  in  the  centre,  retaining 
instead  a  solid  homogeneous  fleshy  character  of  dull  gray- 
yellow  color.  Upon  section  these  have  been  found  whoUy 
cellular  in  construction.  To  distinguish  them  from  the 
ordinary  nodules  they  have  been  designated  tuberculo- 
mata.  Lesions  of  this  kind  may  occur  along  the  lymphatic 
paths,  indeed  seem  more  common  in  the  lateral  cervical 
and  thoracic  chain,  and  upon  bones  and  nerves.  When 
they  are  numerous  the  nodular  caseous  type  is  incon- 
spicuous.,  They  suggest  the  bovine  infection  (Pearl  dis- 
ease), but  one  attempt  to  prove  this  failed.  We  are  of 
the  opinion  that  this  is  the  avian  lymphatic  form,  as 
our  examples  correspond  to  the  literary  descriptions  of 
cervical  tuberculous  lymphatics  in  birds.  No  especial 
variety  of  bird  is  more  often  affected  by  this  process. 

Intestinal  tuberculosis  among  the  Aves  may  be  said  to 
assume  three  forms.  The  best  known,  indeed  the  form 
usually  spoken  of  as  representing  the  common  picture,  is 
that  which  produces  varying  sized  nodules  upon  the 
serosa,  sometimes  associated  with  adhesions  to  neigh- 
boring intestines.  Just  how  this  type  develops  is  not 
known.  In  some  quarters  it  is  believed  to  originate  by 
the  penetration   of  the  tuberculous   granulation  tissue 

33 


506   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

from  the  mucosa  through  the  intestinal  wall  by  following 
lymphatic  channels  and  that  irregular  contractions  of  the 
musculature  squeeze  the  exudate  outward  under  the 
serosa.  Other  observers  think  that  the  bacilli  are  car- 
ried; -1;^  the  lymphatics  to  the  superitoneal  tissue, 
there  starting  the  tubercle.  The  truth  of  the  matter  will 
probably  be  that  both  methods  are  operative  although  we 
have  seen  more  cases  suggestive  of  the  second  than  of  the 
first  explanation.  When  these  peritoneal  nodules  are 
numerous  and  prominent,  mucosal  ulcers  are  uncommon 
and  vice  versa. 

The  second  form  is  the  ulcerative,  flat  ragged  or 
crateriform  defect  situated  in  a  diffusely  thickened  wall. 
This  was  well  illustrated  in  cases  of  pulmonary  infection 
in  doves  and  guans,  suggesting  reinfection  of  the  gut 
tract  from  swallowed  tubercle  bacilli  or  a  backward 
development  of  the  disease  after  the  lungs  were 
nearly  solid. 

The  third  form  of  tuberculous  enteritis  is  quite  inter- 
esting and  striking.  It  is  best  seen  in  the  duodenal  loop 
but  may  occur  anywhere.  Diffuse  thickening  of  the 
enteric  wall  is  noted,  and  when  palpation  is  practiced  a 
resilient  but  leathery  sensation  is  obtained.  Careful 
inspection  reveals  the  mucous  surface  to  be  velvety,  a  con- 
dition due  to  a  swelling,  that  is  widening,  of  the  villi  which 
retain  their  erect  position  and,  when  washed  in  flomng 
water,  will  be  seen  to  move  like  a  field  of  grain  in  a 
breeze.  The  serosa  may  be,  usually  is,  negative.  Studied 
microscopicaUy  the  peculiarity  of  this  form  is  in  the 
development  of  tubercles  and  diffuse  cellular  exudation  in 
the  villus  stalk,  sometimes  extending  into  the  submucosa 
also.  Round  cell  masses  like  lymph  follicles  are  some- 
times prominent.  This  form  is  not  associated  with  any 
peculiar  organic  distribution  so  far  as  my  studies  go. 

Passeres.  The  peculiarity  of  this  order  seems  to  be  in 
the  predominance  of  the  pulmonary  route  as  origin  of 
tuberculosis.    Perhaps  in  no  other  order  has  there  been 


Fig.   54. — TUBERCULOUS  MASSES  OF  INTESTINAL  WALL  SHOWING  OUTWARD 
GROWTH.     SOMETIMES  THESE  MASSES   OBSTRUCT  THE  LUMEN. 


THE  COMMUNICABLE  DISEASES  507 

sucli  extensive  and  advanced  lesions  as  in  these  little 
birds.  Sometimes  one  whole  lung  will  be  solid  while  its 
fellow  will  be  half  occupied  by  caseous  material.  The 
doves  alone  seem  to  approximate  the  Passeres  in  ability 
to  live  with  so  much  tuberculous  exudate. 

Picarise.  Specimens  from  this  order  illustrate  well  the 
intestinal  origin  and  distribution  of  tuberculosis.  There 
was,  among  these  birds,  one  case  sho\ving  tuberculomata 
which  was,  because  of  its  gross  anatomy,  listed  as  the 
pearl  type.    Its  description  is  as  follows : 

Lesson's  Motmot  (Momotus  lessoni).  The  region  above  and  behind 
the  right  clavicle  in  front  of  the  brachial  plexus  on  the  internal  sur- 
face of  the  thorax,  exterior  to  the  first  and  second  ribs,  and  on  the 
internal  surface  of  the  ribs  at  the  junctions  of  ribs  with  the  alae  of 
the  sternum,  there  are  many  small,  irregular,  smooth,  firm,  yellowish 
Avhite  nodules  varj-ing  in  shape  from  spherical  to  sweet  potato  and 
in  size  from  3x3  mm.  to  3  x  7  mm.  These  are  found  quite  homogeneous 
on  cross  section.  They  do  not  resemble  tubercle  or  mould  infection 
but  make  one  think  of  neuromata.  There  are  also  a  few  present  in 
the  left  lateral  air  sacs,  close  to  but  not  joining  the  intestine.  The 
lungs  are  apparently  normal.  Histological  section  of  the  masses  de- 
scribed as  distributed  along  the  nei-ves  consist  of  sharply  outlined  but 
not  well  encapsulated  masses  made  up  of  irregularly  disposed  bunches 
of  large  cells  with  vesicular  nuclei  in  a  stroma  of  loose  connective  tis- 
sue very  inconspicuous  in  amount.  There  is  also  quite  a  number  of 
small  round  cells  and  a  few  leucocytes.  The  large  cells  first  described 
have  the  nucleus  eccentric  for  the  most  part.  Many  of  them  have 
two  nuclei  and  a  few  three  and  occasionally  a  giant  cell  is  observed. 
Blood  vessels  have  a  very  delicate  wall  and  are  frequently  encountered 
in  the  centre  of  these  masses.  Atypical  mitoses  can  be  found.  Here 
and  there  a  seal  ring  placement  of  the  nucleus  can  be  found.  A  few 
eosinophiles  are  present  not  definitely  placed.  Necroses,  with  large 
quantities  of  nuclear  fragments,  are  scattered  irregularly  through  the 
mass.  The  diagnosis  rests  between  an  infectious  granuloma,  false 
neuroma  and  sarcoma.  Tubercle  bacilli  were  found  by  stain  in  great 
numbers  both  within  and  without  the  cells. 

Psittaci.  Tuberculosis  occurs  in  this  order  somewhat 
more  frequently  in  the  varieties  whose  habitat  is  the 
Eastern  world,  although  South  American  birds  also  suffer 
from  it  in  the  characteristic  manner.  There  seems  to  be 
no  difference  in  the  pathology  of  these  two  groups.  Par- 
rots present  very  beautifully  the  separate  solid  or  semi- 


508   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

solid  nodules  of  avian  tuberculosis,  whether  they  be  in 
the  lungs,  liver,  or  spleen.  "VYlien  the  lung  becomes  rid- 
dled with  masses,  coalescence  occurs  and  the  whole  mass 
turns  into  a  cast  of  the  hemithorax.  Lesions  in  the  liver 
are  mostly  isolated,  but  the  spleen  often  appears  like  one 
large  pink  tuberculous  nodule.  The  liver  occupies  as 
usual  the  first  place  in  organic  incidence. 

Striges.  Owls  (and  Struthiones — see  below)  present 
the  interesting  exception  to  the  rule  of  intestinal  origin 
of  tuberculosis  in  birds.  Perhaps  the  platting  is  incorrect 
but  the  birds  in  the  order  under  discussion  had  older  and 
much  more  advanced  lesions  in  the  lungs  and  thoracic 
serosa  than  they  did  in  the  abdominal  organs.  That  this 
was  true  in  all  three  examples  is  in  itself  noteworthy. 
Perhaps  they  possess  less  pulmonary  and  more  intestinal 
resistance.  One  of  these  birds  showed  a  small  recent 
cavitation  in  the  posteroinferior  angle  of  one  lung. 

Accipitres.  With  one  exception  the  cases  of  this 
order  occurred  among  the  Falconidse,  that  is  in  hawks, 
buzzards,  and  eagles.  Their  lesions  are  usually  gen- 
eralized as  indicated  by  the  figures  for  visceral  distribu- 
tion, but  that  half  the  number  should  have  the  oldest, 
most  prominent  lesions  in  the  lungs  is  curious.  Their 
intestinal  tuberculosis  seems  mostly  of  the  diffuse  infil- 
trative type.^ 

Columbae.  These  birds  are  obviously  the  most  sus- 
ceptible of  all  the  varieties  of  which  there  are  sufficient 
autopsies  to  make  a  comparison.  Generalized  nodular 
lesions  emanating  from  the  intestinal  tract  comprise 
their  usual  form,  while  most  of  the  hepatic  lesions  are 
small  miliary  and  nodular;  occasionally  one  sees  caseous 
masses  destroying  large  sections  of  the  organ.  Their 
intestinal  lesions  may  assume  any  of  the  three 
forms  described. 

Galli.  This  is  an  order  of  something  over  the  average 
percentage  incidence  for  the  birds  but  containing  families 
that  seem  very  susceptible  to  tuberculosis.;    The  small 


THE  COMMUNICABLE  DISEASES  509 

immber  of  Brush  Turkeys  (Catheturus  lathami)  had  60 
per  cent,  of  the  disease,  while  South  American  Cracidae 
had  44  per  cent.  These  two  groups  raise  the  incidence  for 
the  order.  Galli  as  a  group  have  generalized  nodular 
tuberculosis  originating  by  the  intestinal  route.  This  is 
especially  seen  in  the  Phasianidae,  while  the  very  sus- 
ceptible Cracidae  have  much  more  prominent  lesions  in 
the  lungs,  often  of  a  massive  caseous  type.  It  is  really 
astonishing  at  times  how  much  of  the  pulmonary  tissue 
is  occupied  by  infiltrate  before  death  has  supervened.  » 
Fuhcarias  are  represented  by  a  special  contingent 
of  rails  and  gallinules.  Avian  characters  are  well  illus- 
trated in  the  order.  So  too  the  succeeding  order,  Alector- 
ides,  another  variety  of  shore  birds,  run  true  to  the  avian 
form.  It  is  interesting  to  note  that  in  the  two  cases 
from  each  of  these  orders  tuberculosis  and  aspergillosis 
have  been  combined.  The  former  has  assumed  the  firm 
nodular  type,  while  the  mycosis  has  been  of  the  air  sac 
variety.  The  following  case  is  worth  citing  as  possibly 
illustrating  infection  per  cloacam.  There  is,  however, 
no  trace  of  this  bird  having  been  with  a  male  with 
the  disease. 

Demoiselle  Crane  {Anthropoides  virgo)  $.  General  tuberculosis 
including  the  oviduct.  All  organs  are  thickly  beset  by  caseous  tuber- 
culous nodules  except  the  lungs  which  have  only  a  few  scattered  ones. 
The  oviduct  is,  for  its  lower  two-thirds,  much  enlarged,  firm,  tough,  pale 
yellow,  thickly  beset  with  caseous  nodules;  upper  parts  uninvolved. 
The  kidneys  are  definitely  enlarged,  in-egular,  almost  mulberry-like, 
brownish  yellow,  firm  and  tough.  On  section  the  lobules  are  irregular, 
connective  tissue  increased,  urates  in  pelves.  Tubercles  in  intestines 
seem  to  be  wholly  peritoneal.  This  seems  like  a  tuberculosis  of  genital 
origin  judging  from  condensation  of  tubercles  in  the  lower  abdomen 
(mass  around  cloaca).  The  ovaries  are  not  involved.  Lungs  and  tho- 
racic air  sacs  relatively  free.  The  pericardium  shows  a  whitish  thicken- 
ing of  both  layers  due  to  the  presence  of  whitish  granules  like  urates. 
Histological  section  of  kidney  shows  the  capsule  not  greatly  altered. 
Glomeruli  largely  negative  but  a  few  show  hyaline  capsular  thickening 
of  vacuoles  in  tufts  or  fibrosis  in  tufts  or  obliteration  of  whole  structure. 
Tubules  largely  degenerated,  distended  or  distorted.  Interstitial  tissue 
between  the  tubules  definitely  but  irregularly  increased.    No  real  attempt 


510  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

at  regeneration.     Few  vessels  show  perivascular  fibrous  change.     One 
tubercle  seen. 

Anseres.  These  birds  present  no  especial  features  so 
far  as  percentage  or  organic  incidence  are  concerned. 
The  individuals  are  mostly  geese  and  swans,  ducks  being 
somewhat  more  often  affected  by  mycosis  than  by  tubercu- 
losis. However,  both  these  diseases  tend  to  assume  the 
nodular  type  in  Anseres  so  that  the  diagnosis  should  be 
supported  by  bacteriological  discovery  of  the  respec- 
tive organisms. 

Struthiones.  The  marked  feature  of  this  order  is 
the  prominence  of  the  isolated  and  confluent  nodules 
in  the  lungs,  of  apparently  greater  age,  certainly  of  great 
size,  than  similar  lesions  in  the  abdominal  viscera.  Case- 
ation of  the  avian  variety  is  well  illustrated  in  these 
birds.  The  thyroid  body  was  involved  in  two  of  the  three 
cases,  the  ovary  in  one.  The  representatives  of  the 
Crypturi,  two  tinamous,  came  at  the  same  time  and  lived 
only  a  few  months.  Miliary  tuberculosis  of  the  small  pre- 
caseous  variety  was  the  form  exhibited  by  both  specimens.*) 

Histology  of  the  Tuberculous  Lesions. 

The  initial  and  characteristic  unit  of  tuberculosis,  the 
miliary  tubercle,  seems  to  be  constnicted  upon  the  same 
general  principles  in  all  cases  of  the  disease  and  in 
all  members  of  the  zoological  groups  in  our  study  and  in 
a  manner  entirely  comparable  to  that  well  known  for  man 
and  for  the  domestic  animals.  There  are,  however,  cer- 
tain minute  differences  which  are  interesting  and  may  at 
some  time  become  important.  It  is  customary  to  speak 
of  the  bovine  tubercle  and  of  the  human  variety,  but  there 
are  also  slight  variations  of  the  microanatomy  of  each  of 
these,  while  one  may  find  on  occasion  a  tubercle  of  the 
human  type  in  a  cow  and  vice  versa.  Not  all  the  domestic 
animals  show  the  bovine  form,  although  in  sheep  and 
swine  it  is  approximated  very  closely.  In  the  horse  there 
is  much  greater  tendency  to   a  central  softening  and 


Fig.   55. 
DIFFERENT  VARIETIES  OF  THE  MILIARY  TUBERCLE. 


M 


:j^^' 


«• 


^*<Kj. 


^.'' 


^?4 


?^; 


\<' 


A. — THE  BOVINE  FORM  SHOWING  THE  NUMEROUS  LANGHANS'  GIANT  CELLS,  THE  ABUNDANT 
SMALL  EPITHELIOID  CELLS,  THE  MODERATE  NUMBER  OF  SMALL  ROUND  CELLS,  AND  THE  ACCOM- 
PANYING  CONNECTIVE  TISSUE  INCREASE.     THERE  IS   MODERATE  CASEATION. 


DIFKKRKNI    VARl 


)r    I'HK   MILIARY    1  I  HKRCI.E. 


:-?i'ri 


'Vi^l 


vfi 


,••> 


'^'^ 


^•-«. 


•'« 


*>«  A   'i    *  * 


t  %r'.^ 


'O: 


■i'J] 


•f*: 


^%«i: 


V/fef/t 


-.i-*- '*?'.:•**?'. 


.••  •^   Kr*f,  'f  i"  * » **  J*  ?*  ■- 


-  i-:o 


,^^ 


fi'/» 


/.A: 


:<4 


/:• 


?^a: 


"'  to*?:.*^ 


^^-'^^tV 


:'/o»f-r 


•  * 


.^^ 


H.— THK    HUMAN    TUBERCLE    WITH   CENTRAL   COMPLETE    NECROSLS,    TYPICAL    CJIANT    CELLS.    ABUN- 
UANT  EPIIHELIOIl)   CELLS  AND  THE   RELATIVELY  NARROW  SMALL   ROUND    CELL   MANTLE. 


Fig.   55. 
DIFFKRF.NT  VARIKTIES  OF  THK  .MILIARY  TUBERCLE. 


D. — AN  AVIAN  TUBERCLE  WITH  CENTRAL  SHARPLY  MARGINATED  NECROSIS  CONTAINING 
MUCH  CHROMATIN  DEBRIS,  THE  IRREGULARLY  ARRANGED  POLYNUCLEAR  CELLS  TYPICAL  OF  AVIAN 
TUBERCLES.  THE  SMALL  NUMBER  OF  REGULARLY  ARRANGED  EPITHELIOID  CELLS.  THE  PAUCITY 
OF  SMALL  ROUND  CELLS  AND  THE   PRONOUNCED   CONNECTIVE  TISSUE  MANTLE. 


ABLED 


■  d  orders,  but  it  lias  not  i^ 
1.    j.-uvery.    It  is,  however,  possibn 
'»e  commonly  found  in  monkeys  with  that 
ulates  and  also  to 
m  tuberelo  that  ii 
iguished    from    mammali;  iilosis    ; 

riV'ian  mycosis. 

The  tubercle  of  the  Prim; 
affair 
clos'^ 
ti- 


the large  pale  t 

I    .,,_r.K-       , , 


cliromai.c 
destroyed   ; 
assume  the  den 
Giant  cells  of  tiu 
often  entirely  mis^ 
There  may  be  L  " 
epithelioid  cells 

illg  JlUrii;!,  • 

Tiot   111-.,   n,, 


512   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

The  avian  tubercle  as  it  rests  in  the  tissue  seems  like  a 
sharply  outlined  almost  encapsulated  body.  This  is  in 
part  due  to  the  homogeneity  of  its  structure  and  in  part 
to  the  fibrocellular  condensation  around  the  caseous  part 
of  the  growths.  Examined  from  without  inward,  there  is 
a  round  cell  mantle,  between  the  elements  of  which  course 
fine  but  easily  perceptible  fibrils ;  elastic  tissue  has  been 
seen  among  them.  The  small  cells  continue  more  deeply 
than  the  fibres,  to  be  succeeded  in  prominence  by  epithe- 
lioid cells  of  rather  dense  character,  the  nuclei  especially 
seeming  quite  rich  in  chromatin  and  round.  The 
rotundity  of  the  nuclei  remains  not  only  in  the  single 
separate  cells  forming  the  middle  zone  of  the  cellular 
cortex,  but  can  be  found  in  the  nuclei  of  the  compound  or 
giant  cells  which  comprise  the  internal  layer  lying  upon 
the  necrotic  centre.  These  giant  cells  are  characteristic 
for  the  avian  tubercle  in  assuming  a  form  like  syncytia 
with  nuclei  arranged  in  irregular  radiating  columns.  This 
internal  large  cell  area  may  completely  surround  the 
central  necrosis  or  it  may  be  interrupted  by  the  large 
single  cells.  Tubercle  bacilli  are  more  common  in  and 
between  single  cells  than  multinuclear  ones.  Within  the 
cellular  zone  lies  the  necrotic  centre,  often,  indeed  usually, 
full  of  chromatic  debris.  This  centre  is  commonly  quite 
amorphous  but  occasionally  one  will  see  what  is  probably 
the  remains  of  a  coarse  coagulum.  Between  the  necrosis 
and  the  cells  one  usually  finds  a  split,  a  sort  of  separation 
of  the  gangrenous  from  the  living  part.  Old  tubercles 
with  denser  fibrous  capsule  retain  this  microanatomy 
in  part,  but  the  cellular  zone  gradually  becomes  thinner 
and  thinner  until  all  that  remains  is  a  narrow  cortex  of 
round  cells  and  imperfectly  retained  multinuclear  cells.) 

Tuberculous  granulation  tissue  without  definite  milia 
consists  entirely  of  the  round  cells  with  small  vacuoles 
and  a  fine  but  definite  fibrosis.  Tuberculomata  consist  of 
cells  of  varying  sizes  with  small  round  nuclei.  Interstitial 
fibrosis  is  delicate  and  barely  visible  unless  especially 


W^\;i^l\ 


K-? 


55 —PHOTOGRAPH  OF  YOUNGEST  AVIAN  TUBERCLE  AFTER 
COMPLETE  FORMATION. 


Fig.  57. — TUBKRCULOMA.  .\  SOLID  IL'MOR-I.IKK  MASS,  CONSISTING  OK  CLOSELY  PACKKI) 
LARf;E  CELLS  KITTED  WITH  RKLAIIVELV  SMALL  ROUND  LOOSE  NUCLEL  THESE  CELLS  ARE 
CROWDED  WITH   BACILLL 


THE  COMMUNICABLE  DISEASES  513 

sought.  Giant  cells  may  be  encountered  but  are  not  so 
large  as  in  milia.  Necrosis  occurs  but  not  in  an  orderly 
manner  in  relation  to  cells  as  in  an  isolated  tubercle. 
Tubercle  bacilli  are  very  numerous. 

Types  of  Bacillary  Infection. 

According  to  experimental  and  statistical  research, 
all  the  tubercle  bacilli  of  the  higher  vertebrate  classes  can 
be  infective  for  any  member  of  these  classes.  Thus,  for 
example,  human  bacilli  have  been  found  in  many  orders 
of  mammalia  and  in  birds.  The  bovine  form  has  been 
found  in  swine.  The  lesson  from  this  is  that  while  the 
special  predilection  of  a  variety  of  the  tubercle  bacillus 
may  be  for  one  kind  of  animal,  it  is  potentially  a  vinis  for 
other  kinds.  Hygienic  principles  have  therefore  been  laid 
down  at  the  Garden  which  aim  at  the  protection  of  all 
specimens  from  every  variety  of  tubercle  bacillus.  For 
this  reason  and  because  the  laboratory  has  not  attempted 
extensive  research  on  bacteriology,  few  type  determina- 
tions have  been  made  and  those  at  hand  oifer  nothing  new 
or  unusual;  they  are  noted  here  as  a  matter  of  record. 
Bovine  bacilli  have  been  judged  by  their  slow  growth 
and  inf ectivity  for  rabbits,  human  bacilli  by  the  reverse 
of  these  characters.  Avian  tubercle  bacilli  can  be  culti- 
vated with  reasonable  ease  directly  from  lesions  not 
bearing  a  mixed  bacterial  flora,  and  grow  in  a  yellow, 
moist,  even,  spreading  colonization.  In  our  two  attempts 
at  infection  of  guinea-pigs,  no  success  was  had,  although 
Rabinowitsch  and  others  had  no  difficulty  in  so  doing ;  this 
strain  may  vary  in  virulence  as  do  other  tubercle  bacilU. 
No  avian  culture  was  obtained  from  a  mammal,  but  a 
bovine  was  found  in  a  parrot  and  a  human  in  a  duck. 
Bovine  bacilli  were  isolated  once  from  a  monkey  (see 
page  496)  and  in  another  case  of  lymphatic  type,  bacilli 
of  the  short  heavy  blunt  shape,  supposed  to  be  character- 
istic of  this  variety  of  the  germ,  could  be  stained.  Monkey 
tuberculosis  in  our  experience  is  usually  due  to  the  human 


514   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

tubercle  bacillus,  judging  by  the  staining  characters  and 
two  successful  cultures. 

Discovery  of  Tuberculosis  During  Life. 

Fully  developed  chronic  tuberculosis  may  be  recog- 
nized with  reasonable  ease  in  the  human  being  and  some 
domestic  animals.  The  diagnosis  rests  largely  upon  the 
history  and  symptoms  and  partly  upon  the  appearance  of 
the  individual  and  upon  signs  elicited  by  physical  exam- 
ination. There  is  good  reason  to  believe  that  these  latter 
methods  are  entirely  applicable  to  certain  wild  animals, 
notably  those  that  can  be  caught  and  held  quiet,  but 
because  of  their  naturally  great  reserve  many  specimens 
offer  little  reason  for  suspicion  as  to  their  tuberculous 
condition  until  near  death.  Certain  ungulates  with 
chronic  pulmonary  disease  get  thin  and  weak  but  remain 
on  their  feet  with  good  appetite  and  satisfactory  dis- 
charges for  many  months.  Primates,  Camivora,  Roden- 
tia  and  Aves  not  uncommonly  come  to  autopsy  with  very 
good  coats  and  without  great  emaciation  and  yet  are 
heavily  infected.  It  can  be  stated  with  fair  positiveness 
that  no  chain  of  historical  data  or  gross  observations  are 
certainly  known  to  us  as  indicative  of  tuberculosis  in  the 
wild  beast.  Coughing  is  not  necessarily  characteristic  of 
chronic  pulmonary  infection,  although  when  continuous  it 
rouses  considerable  suspicion,  especially  in  the  Ungulata. 
It  is  to  be  interpreted  with  care  in  all  animals  that  have 
loose  bedding  as  bits  of  straw  or  seeds  get  into  the  throat 
causing  irritation;  the  dust  of  hay  may  cause  coughing 
in  horses. 

However  much  chronic  or  fatal  tuberculosis  may  be 
interesting  from  the  standpoint  of  pathology  or  of 
zoological  or  visceral  incidence,  the  most  important  fac- 
tors in  our  knowledge  of  the  disease  are  its  early  recogni- 
tion and  treatment,  either  for  curative  or  hygienic 
purposes.  Since  we  have  learned  that  advanced  lesions 
may  exist  in  an  animal  without  materially  affecting  its 


THE  COMMUNICABLE  DISEASES  515 

external  appearance  and  behavior,  it  naturally  follows 
that  early  cases,  possibly  of  an  ''open"  or  infectious 
character  are  still  less  likely  to  give  evidence  of  their 
existence.  This  is  well  recognized  by  veterinarians  as 
being  true  of  cattle,  but  is  perhaps  less  well  known,  or 
possibly  admitted,  by  those  who  handle  the  very  sus- 
ceptible monkey. 

Upon  a  visit  to  a  foreign  garden  I  was  told  that 
experience  alone  is  suflBcient  to  enable  an  observer  to 
detect  tuberculosis,  and  that  the  disturbance  entailed  in 
physical  examination  and  tuberculin  tests  is  preju- 
dicial to  the  well  being  of  all  varieties,  but  especially  the 
delicate  ones.  I  learned  later  that  they  had  the  disease  in 
their  exhibition  cages  all  the  time  but  decided  to  put  their 
method  to  the  test.  Shortly  after  my  return  from  abroad 
a  splendid  specimen  of  Grivet  Monkey  {Cercopithecus 
sah(Bus)  was  condemned  by  the  tuberculin  test.  He  was 
well  studied  by  the  superintendent  and  two  very  experi- 
enced keepers,  all  of  whom  pronounced  him  one  of  the 
finest  specimens  they  had  ever  seen,  and  stated  tliat  he 
was  behaving  quite  normally.  Despite  their  protests  he 
was  sacrificed,  tuberculosis  with  early  cavitation  being 
found  in  the  upper  lobe  of  the  left  lung.  Incidentally 
vague  physical  signs  were  found  by  auscultation,  but  as 
the  monkey  was  unruly  and  had  long  pectoral  hairs  little 
weight  was  placed  on  the  observation.  However,  it  is 
frequently  possible  to  make  very  thorough  physical 
examination  of  the  lungs  of  the  more  tractable  specimens, 
diagnoses  of  pneumonia  and  bronchitis  being  frequently 
made  in  this  and  other  parks,  so  that  treatment  may 
be  instituted. 

Importance  of  Transmission  and  Known  Susceptibility. 
Some  light  upon  possible  reasons  for  the  poor  condi- 
tion of  an  individual  animal  is  of  course  shed  by  a 
knowledge  of  the  disease  to  which  that  particular  variety 
is  most  susceptible,  to  which  may  be  added  the  data 


516   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

obtained  from  previous  deaths  in  the  same  group  or 
enclosure.  Thus,  for  example,  a  sickly  monkey  would  be 
suspected  of  having  tuberculosis  or  early  osteomalacia, 
whereas  no  suspicion  of  these  diseases  would  fall  upon 
the  marsupials.  The  same  position  would  be  assumed  if  a 
dove  and  a  heron  were  out  of  condition. 

In  so  far  as  enclosures  are  concerned,  the  matter  is 
somewhat  different.  "WTienever  a  case  of  tuberculosis 
occurs  in  a  cage,  the  remaining  specimens  if  any  are 
removed  and  the  place  cleaned  by  soap  and  water  and 
disinfectant  and  paint.  The  naked  flame  from  a  blast 
lamp  is  used  when  possible.  Out-of-doors  enclosures  are 
vacated,  spread  with  lime  and  allowed  to  lie  fallow  for  as 
long  a  time  as  practicable.  These  methods  have  been  in  the 
main  successful  in  clearing  a  cage  of  the  disease,  and  all 
our  experience  demonstrates  the  effect  of  cage  hygiene 
and  the  selection  of  non-infective  replacements.  A  few 
places  such  as  those  occupied  by  doves  and  guans  have 
not  been  freed  of  infection,  if  one  judge  by  its  appearance 
when  new  specimens  are  placed  in  them,  but  they  may  of 
course  be  due  to  the  infection  from  elsewhere.  The  his- 
tory of  seven  years  in  the  new  bird  house  where  the 
hygienic  conditions  are  excellent,  seems  to  indicate  that 
a  cage  thoroughly  cleaned  is  no  longer  a  source  of  danger, 
and  that  a  repetition  of  tuberculosis  in  such  an  enclosure 
is  due  to  its  importation  with  new  exhibits. 

The  spread  of  the  disease  to  nearby  cages  seems  to 
depend  upon  two  factors.  If  the  number  of  cases  has 
been  large  and  the  infection  virulent,  immediately  adjoin- 
ing cages  are  involved,  but  the  tendency  to  spread  is 
directly  proportional  to  the  proximity  of  orders  or 
families  that  have  a  high  susceptibility  for  tuberculosis. 
This  second  factor  seems  to  be  the  more  important  and 
is  illustrated  by  our  experience  in  one  corner  of  the  new 
bird  house.  In  this  area  are  exhibited  certain  doves  and 
pheasants,  among  which  are  many  cases,  while  the  pas- 
serine varieties  nearby  are  little  affected.    So  too  in  the 


THE  COMMUNICABLE  DISEASES  517 

flying  cage  the  disease  has  occurred  in  varieties  with  high 
general  susceptibility.  There  are  at  the  present  writing 
ten  orders  on  exhibition  in  this  large  enclosure  and  there 
have  been  more.  An  occasional  case  of  the  disease  occurs, 
but  only  in  the  orders  which  show  it  elsewhere.  The 
Herodiones,  of  which  we  have  had  nearly  one  hundred 
autopsies  and  many  now  are  on  exhibition,  are  always 
well  represented  in  this  cage  and  yet  show  no  tubercu- 
losis. In  the  ten  orders  mentioned  above  three  show  no 
cases  of  the  disease. 

These  observations  illustrate  the  spread  of 
tuberculosis,  especially  to  the  most  susceptible  varieties, 
and  how  non-susceptibles  under  good  hygienic  conditions 
fail  to  become  infected  even  when  infected  animals  are 
near  them.  The  freedom  of  activity  in  the  large 
enclosure  is  doubtless  an  important  factor. 

The  history  of  the  past  three  years  with  regard  to  the 
control  of  tuberculosis  in  the  small  cages  shows  that 
twenty-nine  were  infected,  but  by  the  measures  employed 
nineteen  have  remained  free  of  the  disease  for  one  year ; 
three  of  the  remaining  ten  are  known  to  have  received 
newly  arrived  and  possibly  infected  specimens. 

The  accredited  method  of  transmission  in  birds,  the 
swallowing  of  material  soiled  with  the  feces  richly  laden 
with  germs,  is  the  principal  reason  why  infected  en- 
closures and  their  immediate  environment  are  the  prin- 
cipal breeding  places  for  tuberculosis.  To  be  sure  air 
currents  may  blow  the  virus  around,  allowing  it  to  light 
upon  food  in  other  cages  but  this  cannot  be  a  great  menace 
if  for  no  other  reason  than  that  we  have  had  no  epizootic 
outbreak  of  the  disease,  when  there  were  groups  of  deaths 
in  doves  and  guans. 

Evidences  with  which  to  trace  transmissions  are  much 
clearer  in  the  birds  than  in  the  mammals  with  the  excep- 
tion of  monkeys  and  some  ungulates.  Of  course  cases 
are  perhaps  too  few  in  the  carnivores  and  rodents  to 
permit  correct  deductions  but  it  is  very  rare  that  more 


518   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

than  one  case  occurs  in  the  same  enclosure  containing 
groups  of  these  varieties.  Nor  do  animals  in  adjoining 
cages  seem  to  ''  catch  "  the  infection.  This  observ^ation 
does  not  suggest  that  any  relaxation  of  hygiene  need  be 
allowed  but  probably  it  implies  that  not  many  bacilli  are 
excreted  by  these  animals;  they  cough  very  rarely. 
Groups  of  ungulates  (bison  and  deer)  are  often  known 
to  be  infected  but  just  how  it  has  arisen  is  seldom  clear. 
Transmission  from  monkey  to  monlvey  has  been  observed 
so  frequently  that  it  cannot  be  doubted,  nor  mil  anyone 
wonder  at  it  if  reflection  is  given  to  the  close  personal 
contact  of  these  animals  during  their  natural  behavior. 
They  huddle,  pluck  lice  from  one  another,  take  food 
from  the  mouth  of  another,  bite  and  perform  many  other 
actions  greatly  facilitating  the  transfer  of  any  virus. 
Bacilli  may  also  be  disseminated  by  cougliing,  drooling 
and  with  the  fecal  discharges,  for  which  latter  there  seems 
ample  opportunity  since  a  notable  percentage  of  cases 
have  intestinal  lesions.^  Monkeys  do  not  seem  to  raise 
sputum  and  expectorate  it  but  they  do  eject  saliva  from 
their  lips. 

Contraction  of  the  disease  from  infected  cages  is  be- 
lieved to  have  occurred  at  least  once  in  our  experience 
but  the  lesson  of  complete  sanitary  cleaning  of  the 
enclosure  learned  from  that  happening,  seems  to  have 
enabled  us  to  forestall  its  repetition. 

Tjhe  Tuberculin  Test. 

Tuberculosis  presents  the  greatest  single  problem 
among  the  specific  infectious  diseases  which  the  director 
of  a  menagerie  must  attempt  to  solve.  Even  though  one 
may  possess  a  knowledge  of  its  zoological  distribution, 
clinical  characters  and  pathological  effects,  these  are  in- 
sufficient criteria  for  its  detection  at  a  stage  when  the 
animal  might  be  saved  by  treatment  or,  what  is  most 
important,  removed  from  its  companions  that  they  might 
be  protected.     To  this  end  there  remains  but  a  single 


THE  COMMUNICABLE  DISEASES  519 

procedure  for  the  discovery  of  the  existence  of  tubercu- 
losis— the  use  of  tuberculin  in  one  of  its  forms  by  one 
of  its  methods  of  application.  The  use  of  this  test 
in  veterinary  medicine  needs  no  commentary,  having 
made  its  place  in  clinical  and  hygienic  practice  for  a 
quarter  century  or  more.  Armed  with  the  knowledge  of 
the  satisfactory  use  of  the  toxins  of  tubercle  bacillus  in 
cows,  Dr.  Penrose,  Dr.  C.  Y.  White,  Dr.  A.  E.  Brown  and 
Dr.  Leonard  Pearson  began  in  1901  a  series  of  experi- 
ments with  old  tuberculin  of  Koch  which  have  led  to  the 
development  of  a  technique  for  its  use  in  the  detection 
of  infected  monkeys.  These  interesting  and  instructive 
animals,  being  known  as  highly  susceptible  since  most 
of  the  collection  died  of  the  disease  in  those  days,  and 
being  handled  with  reasonable  ease  by  experienced  men, 
were  investigated  as  the  most  important  specimens  upon 
which  to  perfect  the  method.  Other  varieties  have  been 
studied  since  and  I  shall  refer  to  them  individually.  The 
greatest  amount  of  work  and  the  most  conspicuous  suc- 
cess attended  the  observations  upon  monkeys  and  the 
results  of  this  study  are  now  in  daily  use  in  this  Garden. 
The  work,  conclusions  and  results,  originated  by 
Doctor  Penrose,  Doctor  White,  and  Doctor  Brown 
can  be  described  as  one  of  the  most  completely  satis- 
factory series  of  observations  in  scientific  medicine. 
Applying  the  principle  that  a  tuberculous  animal  reacts 
to  the  injection  of  tuberculin  by  a  temperature  rise, 
the  normal  temperature  curve  of  the  monkey  was  studied, 
that  of  the  tuberculous  monkey  determined  by  killing 
many  specimens.  This  enabled  them  to  state  which 
animal  was  infected,  which  was  not  and  to  place  on  exhibi- 
tion only  healthy  specimens.  Added  to  tliis,  strict  hygi- 
enic principles  in  the  housing  and  handling  of  the  animals 
have  resulted  in  the  elimination  of  the  disease  from  our 
exhibition  house.  Occasionally  a  case  may  develop,  per- 
haps from  feeding  by  visitors,  but  the  matter  is  no  longer 
a  problem.    I  know  of  no  more  complete  and  satisfactory 


520   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

experiment  and  its  practical  application  than  this  work, 
which  is  condensed  in  the  succeeding  paragraphs. (2) 

The  Temperature  of  Monkeys. 

The  success  of  the  tuberculin  test  in  the  lower  animals 
as  in  man  depends  chiefly  upon  the  alterations  in  tempera- 
ture following  the  injection  of  the  toxin.  It  is  generally 
admitted  to-day  that  a  healthy  animal's  temperature  will 
not  be  affected  by  the  introduction  of  this  material.  There 
are  in  addition  changes  in  the  pulse  and  respiration  rate 
and  in  the  physical  signs  but  these  are  detected  with 
difficulty  and  are  much  less  definite  than  thermometric 
records.  The  first  essential  was  therefore  a  thorough 
familiarity  with  the  normal  temperature  of  the  monkey, 
a  requirement  which  met  with  considerable  difficulty  from 
the  beginning  since  the  earliest  observations  revealed  puz- 
zling irregularities.  This  necessitated  the  establishment 
of  certain  regulations  of  teclinique  which,  after  the  pre- 
liminary tests,  have  been  found  satisfactory  enough  to 
continue  until  the  present  day.  All  monkeys  are  received 
in  the  quarantine  rooms  of  the  laboratory  where  they  are 
observed  by  the  officials  of  the  Garden  and  of  the  labora- 
tory and  there  they  remain  in  separate  cages  until  passed, 
as  free  from  tuberculosis,  to  the  exhibition  house. 

The  handling  of  monkeys  for  the  purpose  of  taking 
temperatures  is  a  matter  of  no  small  importance  since 
excitement  will  quite  definitely  increase  the  registration. 
We  have  been  fortunate  enough  to  have  in  charge  of  this 
work  since  its  inception  the  same  man,  Keeper  McCrosson, 
who  is  thoroughly  experienced  in  the  care  of  these  beasts 
and  who  can  catch  and  hold  them  with  a  minimum  of  dis- 
turbance. To  him  and  to  the  interested  laboratory  helpers 
much  credit  is  due.  Small  specimens  like  capucins  and 
spider  monkeys  are  caught  with  the  gloved  hand  or  with 
the  protection  of  a  piece  of  heavy  cloth.  Larger  specimens 
may  be  caught  in  a  net  while  strong  monkeys  are  fitted 

(2)  White  and  Fox,  Archives  of  Internal  Medicine,  1909,  Vol.  IV, 
p.  517. 


THE  COMMUNICABLE  DISEASES  521 

with  a  collar  and  chain  by  which  they  are  pulled  into  the 
corner  of  the  cage  and  held,  while  the  door  is  opened  to 
permit  a  helper  to  catch  the  feet  and  arms.  Two  experi- 
enced men  can  take  the  temperature  of  any  monkey  that 
can  be  handled  at  all  safely.  The  knowledge  of  how 
to  do  such  work  reduces  the  excitement  of  the  animal 
and  renders  more  accurate  the  observation  of  its  tempera- 
ture. During  the  period  of  temperature-taldng  food  is 
given  in  small  quantities  and  only  after  the  record  is 
made. 

Temperatures  are  taken  in  all  animals  by  rectum,  (3) 
the  thermometer,  a.  separate  instrument  but  always  the 
same  for  each  animal,  well  greased  with  plain  vaseline, 
being  passed  along  the  anterior  rectal  wall  and  allowed 
to  register  for  twice  its  indicated  speed.  During  the  pre- 
liminary work,  special  instruments  of  officially  standard- 
ized accuracy  were  obtained  by  Doctor  Brown  but  once 
the  normals  were  obtained,  ordinary  good  thermometers 
registering  from  94°  F.  to  108°  F.  have  been  employed. 
If  the  record  vary  very  much  from  the  expected,  such 
as  the  figures  obtained  at  the  same  time  on  the  preceding 
day,  or  if  the  rectum  be  crowded  with  feces,  the  instru- 
ment is  shaken  down  and  reintroduced.  In  order  to 
facilitate  timing  of  exposures  we  use  sand  glasses  of 
three  minute  run. 

After  some  experimentation  by  taking  records  at  vari- 
ous times  of  day  it  was  found  that  monkeys  as  a  group 
do  not  have  a  uniform  temperature  during  twenty-four 
hours  but  register  a  higher  figure  during  daylight  than 
during  darkness.  This  is  probably  due,  as  I  shall  dis- 
cuss, to  the  period  of  activity,  not  to  the  time  of  day. 
Figures  obtained  at  various  hours  indicated  that  the 
highest  and  lowest  temperature  would  be  obtained  if 
records  were  made  at  four-hour  intervals  at  three,  seven 
and  eleven  o  'clock  AM.  and  PM.  To  give  the  normal  tem- 

(3)  The  temperature  in  the  axilla  is  often  .5°  F.  hi^rher  than  by 
rectum,  but  the  difficulties  of  the  axillary  method  render  it  impracticable. 

34 


522   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

perature  of  a  moiikey,  the  kind  and  the  time  of  day  are 
necessary  adjuncts.  Reference  to  forty-eight  hour  charts 
which  are  used  for  the  illustration  of  normal  records,  and 
for  contrast  with  tuberculin  reactions  later,  wdll  convey  to 
the  reader  a  better  idea  of  the  normal  daily  rhythm 


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Chart  A.    ORANG  UTAN  (Simia  satyrus).     Non-tuberoulous  at  death. 

of    the    simian    heat    regulating    system    than    would 
verbal  description. 

The  anthropoid  apes  (Cur\^es  A  and  B)  have  on  the 
whole  a  mean  temperature  nearer  the  human  being  than 
do  the  lower  monkeys,  but  they  too  present  daily  variables 
far  greater  than  man.  The  high  point  of  their  curve,  at 
three  PM.,  is  in  the  neighborhood  of  100°  F.  the  lower 

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ing tuberculin. 


THE  COMMUNICABLE  DISEASES 


523 


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low  point  at  three  AM.,  there  is  in  them  not  by  any  means 
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Chart  B.    CHIMPANZEE  (Pan  niger).     Non-tuberculous  at  death. 

Cebidae.  Our  records  of  temperatures  in  the  Hylobates 
(Gihhons)  are  not  extensive  enough  to  quote  but  what  we 
have  approach  those  of  the  lower  monkeys. 

Graphic  curves  of  the  normal  temperatures  of  the 
various  genera  of  Cercopithecidae  and  Cebidae  present 
striking  similarities  in  the  regularity  with  which  the 
daily  rhythm  is  performed.  In  the  seven  genera  of  which 
we  have  accurate  records  the  normal  high  points  fall 
between  102°-103°  F.  and  the  low  points  between 
99° -100°  F.  while  the  curve  of  the  four-hourly  steps  is 


524   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

closely  comparable.  The  curves  D  to  J  are  composites 
from  charts  of  animals  that  have  been  tested  with  tuber- 
culin, wliich  thereafter  died  or  were  killed  and  found  free 
from  tuberculosis.  Not  every  individual  chart  that  may 
come  to  hand  necessarilv  follows  the  exact  course  detailed 


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Chart  C.    Composite  chart  of  twenty-two  non-tuberculous  Lemures. 

in  these  illustrative  curves  but  these  latter  offer  a  guide 
as  to  what  is  to  be  expected  of  the  different  varieties. 
They  show  unequivocally  the  V-shaped  curve  of  tke 
temperature  of  the  monkey  during  twenty-four  hours. 

The  CallitrichidaB  or  Hapalidae  have  failed  to  show 
tuberculosis  in  our  Garden  and  little  has  been  done  upon 
them.  As  a  matter  of  record  there  is  reproduced  the 
only  satisfactory  chart  at  hand  (K)  taken  very  early  in 


THE  COMMUNICABLE  DISEASES 


525 


the  researches.  It  shows  a  similarity  to  those  of  the 
higher  monkeys ;  because  of  its  very  high  afternoon  record 
the  animal  was  killed ;  no  tuberculosis  was  found. 

The  Lemures,  being  close  to  the  Primates  zoologically 
and  presenting  a  high  incidence  of  tuberculosis,  were 


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Chart  D.     Composite  chart  of  eighteen  non-tuberculous  Cercopithecus. 

included  in  this  study.  Observations  upon  their  normal 
temperature  were  hampered  more  than  upon  that  of 
monkeys  and  even  to-day  we  cannot  feel  the  same  confi- 
dence in  the  records.  Irregularity  is  most  marked  and 
they  seem  easily  disturbed  by  handling.  Chart  C  shows 
a  composite  temperature  for  forty-eight  hours  of  twenty- 
two  proven  non-tuberculous  Lemures.  The  tendency  for 
the  ''  night  drop  "  is  certainly  existent  but  with  much 
less  definiteness  than  in  the  Primates. 


526   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Conditions  Which  Modify  the  Temperature. 

Observations  by  A.  E.  Brown  (4)  and  by  Simpson  and 
Galbraith  (5)  would  seem  to  indicate  that  the  diurnal  vari- 
ation in  monkeys  is  due  to  periodicity  of  activity.  Doctor 
Brouni  found  that  the  temperature  of  a  night  monkey  is 


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Chabt  E.    Composite  chart  of  seventeen  non-tuberculous  Macacus. 

reversed,  that  it  is  higher  during  the  dark  than  the  day- 
light hours.  See  chart  of  Potto  {Perodictus  potto)  chart 
L.  The  Scotch  observers  report  that  if  the  activity  of 
day  monkeys  were  reversed,  daytime  being  made  artifi- 
cially dark  and  activity  forced  during  the  night,  the  tem- 
perature curves  were  likewise  reversed. 

(4)  A.  E.  Brown,  Proc.  London  Zool.  Soc,  June,  1909,  p.  81. 

(5)  Simpson  and  Galbraith,  Trans.  Royal  Soc,  Edinburgh,  XIV,  p.  1, 
65,  1906. 


THE  COMMUNICABLE  DISEASES  527 

Perhaps  the  most  important  discoveries  of  these  inves- 
tigators concerned  the  influence  of  excitement  upon  the 
temperature  records.  These  observers  indicate  definitely 
that  the  greater  the  physical  activity  and  nervous  excite- 
ment the  higher  the  thermometric  record.    We  have  noted 


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Chart  F.    Composite  chart  of  eleven  non-tuberculous  Papio. 

that  the  substitution  of  a  strange  keeper  who  may  not 
be  gentle  and  tactful  with  the  monkeys  can  serve  to  raise 
the  temperature  above  the  records  obtained  by  an  experi- 
enced man  with  whose  methods  the  animals  are  familiar. 
A  knowledge  of  these  facts  dictates  at  least  two  im- 
portant precautions  on  our  part — our  specimens  must  be 
kept  under  identic  conditions  peculiar  to  their  kind,  and 
surroundings  must  be  established  offering  comfort  with 
a  minimum  of  annoyance  in  transfer  and  handling.    To 


528   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

this  end  all  specimens  upon  receipt  are  put  into  separate 
cages  suitable  to  their  size  and  allowed  to  become  accus- 
tomed to  their  surroundings  for  several  days  before 
attempt  at  temperature-taking  is  made.  Mention  has 
already  been  made  of  the  experience  and  interest  of  the 
principal  keeper;  the  regularity  of  records  is  an  attest 
to  his  work.  Daily  three  o'clock  afternoon  temperatures 
are  taken  first  to  accustom  the  animal  to  the  matter,  be- 
fore test  records  or  tuberculin  injections  are  made. 

The  existence  of  pathological  states  undoubtedly 
affects  normal  temperature  curves  and  tuberculin  reac- 
tions. Gastroenteritis  has  the  effect  of  increasing  the 
whole  level  and  of  making  irregular  the  midday  and 
afternoon  records.  Respiratory  tract  disease  cannot  be 
said  to  have  a  very  definite  effect ;  its  most  frequent  in- 
fluence seems  to  be  to  drive  the  night  records  lower  so 
that  there  is  a  long  fall  between  seven  and  eleven  PM.  and 
a  long  rise  between  seven  and  eleven  AM. 

The  Test. 

The  preliminary  rest  of  the  new  arrivals  having  passed 
daily  three  o'clock  afternoon  temperatures  are  taken  un- 
til an  even  level  is  obtained;  this  requires  usually  four 
days  but  in  very  nervous  specimens  it  may  be  much 
longer.  The  afternoon  temperature  course  provides  not 
only  a  means  of  teaching  the  monkey  what  is  coming  but 
supplies  us  with  a  high  point  record  for  comparison. 
When  a  new  variety  is  received,  a  full  normal  twenty-four 
hour  record  is  usually  made.  This  preparatory  routine 
being  fulfilled,  the  animal  is  injected  under  the  skin  of  the 
tliigh  or  flank  with  freshly  diluted  mixed  bo\dne  and 
human  tuberculin. 

The  Dosage. 

Early  trials  with  this  substance  revealed  the  fact  that 
a  dosage  based  upon  the  weight  in  comparison  to  man 
failed  to  elicit  a  definite  response  whereas  if  based  upon 


THE  COMMUNICABLE  DISEASES  529 

relative  weight  of  cow  was  too  large.  The  finally  deter- 
mined quantity  was  arrived  at,  as  was  the  case  in  early 
human  and  bovine  work,  by  experiment  and  trial  and 
was  as  follows:  A  monlcey  of  five  to  ten  pounds  (2.3 
to  4.5  kilos)  received  an  initial  dose  of  1.  milligram 
and  for  each  additional  five  pounds  (2.3  kilos)  0.5 
mg. ;  this  is  0.2  to  0.4  mg.  per  kilo.  The  amount  given 
to  man  varies  from  2.  to  5.  mg. ;  if  the  body  weigh  60  kilos 
this  is  0.03  to  0.08  mg.  per  kilo.  Cows  are  given  usually 
in  this  country  400  mg.  or,  for  a  cow  of  250  kilos,  1.6  mg. 
per  kilo.  In  the  early  work,  doses  comparable  to  the 
figure  for  man  failed,  whereas  at  least  two  animals  died 
very  quickly  after  1.+  mg.  per  kilo ;  5.  mg.  was  the  original 
high  dose.  While  the  death  of  a  monkey  after  a  large 
dose  was  of  no  moment  and  was  perhaps  desirable,  it 
would  only  be  the  heavily  diseased  specimens  and  this 
would  give  no  criterion  upon  which  to  judge  the  appro- 
priate dose  for  all.  Experience  seems  to  warrant  us  in 
continuing  with  our  present  figures  since  all  tuberculous 
monkeys  have  reacted  to  it.  Subsequent  cases  for  retest 
are  increased  from  50  to  100  per  cent,  depending  upon  the 
size  of  the  monkey,  the  very  robust  and  vigorous  ones 
receiving  an  increase  represented  by  the  higher  figure. 
One  monkey  injected  eight  times  has  risen  from  1.  to  24. 
mg.  with  constantly  a  negative  response  over  a  period 
of  nine  years. 

Doses  for  Lemures  are  relatively  higher,  averaging 
1.5  mg.  or  about  0.5  mg.  per  kilo;  they  are  increased  in 
the  same  manner  as  above. 

The  Tempekature  Taking. 

Injections  are  usually  made  in  the  late  forenoon,  tem- 
perature records  being  started  at  the  usual  three  PM. 
hour  and  continued  at  four-hour  intervals  for  forty-eight 
hours,  giving  thirteen  records  over  two  days,  a  time  period 
presenting  two  complete  cycles  of  diurnal  variation.  This 
was  found  necessary  because  certain  cases  do  not  react 


530   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

during  the  first  day.  Explanation  of  this  was  sought 
in  the  nature  of  the  lesion  but  could  not  be  found  further 
than  that  mild  early  lesions  may  give  it  but  it  camiot  be 
read  as  indicative  of  low  activity  since  one  case  of  laryn- 
geal tuberculosis  had  this  "  delayed  reaction."    At  times 


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Chart  G.     Composite  chart  of  five  non-tuberculous  Cynopithecus. 

it  has  seemed  to  occur  when  the  injection  fluid  formed  a 
blister  under  the  skin,  a  pocket  in  the  areolar  subcutane- 
ous tissue,  whence  absorption  would  be  slow.  Whatever 
the  correct  explanation,  experience  has  justified  the  re- 
cording of  temperatures  for  full  forty-eight  hours. 

The  Reaction. 

The  experience  gained  with  these  monkeys  supported 
definitely  the  general  opinion  that  tuberculin  injected  into 
healthy  animals  will  not  disturb  the  temperature  but  will 


THE  COMMUNICABLE  DISEASES  531 

produce  decided  changes  in  that  of  tuberculous  animals. 
The  reaction  in  the  tuberculous  animals  may  assume 
several  characters,  of  which  usually  two  are  combined 
in  a  chart.  The  commonest  and  most  convincing  is  a 
definite  rise  in  the  first  twelve  hours,  amounting  to  one 


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Chart  H.    Composite  chart  of  eleven  non-tuberculou3  Ateles. 

degree  or  more ;  rarely  it  may  be  three  degrees  (W) .  This 
is  followed  either  by  a  maintenance  of  a  liigh  level  or  an 
attempt  to  perform  the  night  drop.  It  may  be  said  that 
in  general  there  is  an  abortive  attempt  in  nearly  all  tuber- 
culin reactions  to  simulate  the  V  of  the  normal  cycle ;  this 
can  be  seen  in  charts  M,  N,  0  and  P.  Another  rise  may 
be  attempted  during  a  similar  period  of  the  second  twenty- 
four  hours  or  the  whole  course  may  at  that  time  approxi- 
mate the  normal.    A  modification  of  this  type  of  reaction 


532   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

is  the  performance  of  the  whole  daily  rhythm  on  a  high 
level,  set,  as  it  were,  by  the  initial  three  PM.  record.  This 
form  is  confusing  at  times  and  has  been  responsible  for  at 
least  one  of  our  mistakes.  Combined  with  this  high  level 
of  curve  is  a  tendency  for  the  second  twenty-four  hours 


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Chart  I.     Composite  chart  of  eight  non-tuberculous  Cercocebus. 

to  be  higher  than  the  first  day  (See  Variegated  Cebus  Q 
and  Sooty  Mangabey  R  and  Chacma  Baboon  W).  The 
second  type  of  reaction,  illustrated  by  chart  S,  fails  to 
resemble  the  normal  daily  cycle  of  the  monkey  tempera- 
ture but  has  sudden  rises  and  falls  as  its  characteristic 
feature.  We  have  learned  to  look  mth  suspicion  on  all 
charts  with  sudden  marked  changes  of  record  even  if  they 
follow  in  the  main  a  rhythmic  course.  The  sudden  fall 
exhibited  by  a  very  sick  monkey  illustrated  by  Grivet 


THE  COMMUNICABLE  DISEASES 


533 


Monkey  (T)  aiid  Campbell's  Monkey  (U)  is  a  bad 
sign.  It  has  been  met  more  often  in  advanced  caseous 
pulmonary  tuberculosis  than  in  any  other  tuberculous 
lesion.  On  two  occasions  it  has  been  seen  in  the  absence 
of  tuberculosis  so  that  retest  is  indicated  if  the  specimen 


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Chart  J.    Composite  chart  of  twenty  non-tuberculous  Cebus. 

be  valuable ;  such  animals  however  rarely  survive  the  dis- 
turbance incident  to  the  test  as  they  are  usually  suffering 
with  some  serious  disease.  Illustrative  charts  of  several 
positive  reactions  serve  to  elucidate  their  character  better 
than  description.  If  comparison  and  contrast  of  the  nor- 
mal and  post-injectional  temperature  be  made,  the  con- 
clusions are  definite. 

It  cannot  be  said  that  any  type  of  reaction  indicates 
a  particular  form  of  disease  although  the  last  type,  the 


534   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

falling  of  the  temperature  beyond  the  thermometric  regis- 
tration point,  usually  means  advanced  lesions  especially 
of  the  caseous  pneumonic  form.  A  very  small  lesion  may 
give  a  definite  reaction  as  in  Cebus  (V). 

The  examples  given  are  those  of  a  definite  character 
but  there  are  many  charts  that  vary  from  the  normal 


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Chart  K.    GEOFFROY'S  MARMOSET  (Leontocebus  geoffroyi).     Non-tuberculous. 

upon  which  a  decision  is  extremely  difficult  to  make.  Such 
animals  are  held  in  quarantine  to  be  retested  after  the 
lapse  of  three  months.  Early  in  the  work  a  suspected 
specimen  was  reinjected  after  two  weeks,  failed  to  give  a 
reaction  but  died  in  about  two  months  of  tuberculosis.  The 
nullification  of  the  test  by  previous  injections  of  tubercu- 
lin is  well  kno^vn.    Three  months '  interval  permits  a  dis- 


THE  COMMUNICABLE  DISEASES 


535 


appearance  of  the  non-sensitivity  and  allows  any  latent 
tuberculosis,  possibly  stimulated  by  the  toxin,  to  develop. 
Eepetition  upon  the  same  monkey  has  occurred  as  high 
as  ten  times  without  apparent  harm. 

There  are  sometimes  in  human  beings  local  reactions 
at  the  point  of  injections.  These  have  been  entirely  lack- 


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Chart  L.    POTTO  (Perodicticus  potto).    Healthy. 

ing  from  our  monkey  specimens.  Nor  have  we  ever  seen 
secondary  tuberculous  lesions  appear  at  the  point  of  the 
needle-stick.  Aseptic  syringes  and  generally  cleanly  tech- 
nique have  also  protected  against  local  abscesses.  When 
an  animal  is  injected  he  may  scratch  or  pick  at  the  spot 
for  a  minute  or  two  but  thereafter  seems  to  ignore  it 
entirely. 


536   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Results. 

The  value  of  the  test  can  best  be  estimated  by  a  recital 
of  the  mortality  of  monkeys,  from  tuberculosis,  since  its 
inception.  Before  the  test  was  started  practically  every 
monkey  in  the  collection  for  sufficient  length  of  time  to 


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Chart  M.     MONGOOSE  LEMUR  (Lemur  mongoz).    Tuberculous. 

be  exposed  died  from  the  disease.  The  average  duration 
of  exhibition  life  of  all  specimens  up  to  1903  did  not 
exceed  eleven  months.  The  time  has  risen  almost  unin- 
terruptedly until  now  it  is  thirty-five  months.  There  are, 
at  time  of  writing,  sixty-eight  specimens  in  the  cages 
which  have  been  on  view  from  one  to  one  hundred  and 
eighty-five  months  with  an  average  of  fifty-four  months. 
These  figures  speak  for  themselves  as  evidence  of  the  re- 
Tlie  average  mortality  from  enteri- 


duction  of  infection. 


THE  COMMUNICABLE  DISEASES 


537 


tis  and  degenerative  bone  disease  has  remained  about  the 
same  through  all  these  years.  Percentage  figures  such  as 
are  recorded  in  our  yearly  report  are  misleading  because 
all  monkeys  written  into  the  property  record  of  the  Gar- 
den are  listed  and  since  some  of  these  specimens  remain  in 


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Chart  N.    BLACK-HANDED  SPIDER  MONKEY  (Ateles  geoffroyi).    Tuberculous. 

quarantine,  they  do  not  properly  belong  to  the  exhibition 
collection.  Up  to  1906  when  the  test  technique  was  perfec- 
ted nearly  all  deaths  were  due  to  tuberculosis,  the  figure 
for  1906  (including  experimental  animals)  being  78  per 
cent.  However  from  February  1906  to  October  1907  and 
from  then  until  May  1910  no  case  of  tuberculosis  occurred 
in  the  exhibition  cages  and  both  deaths  at  these  given 
times  seem  like  infection  from  ^dsitors.  During  the  next 
three  years  thirteen  monkeys  died  of  the  disease  in  the  ex- 

35 


538  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

hibition  and  many  more  in  quarantine.  By  1913  the  out- 
break was  stamped  out.  Its  explanation  is  not  so  very  far 
to  seek.  In  the  fall  of  1910  we  obtained  some  suspected 
monkeys  which  were  kept  in  one  of  the  quarantine  rooms. 
After  repeated  testing  two  were  passed.  From  them  five 


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Chart  O.    RHESUS  MACAQUE  (Macacus  rhesus).    Tuberculous. 

cases  are  known  to  have  originated  and  it  was  not  until  in 
1912  when  the  whole  exhibition  house  was  cleaned  of  speci- 
mens, thoroughly  disinfected  and  fumigated  and  until 
every  specimen  was  retested,  that  the  infection  passed. 
In  1914  no  cases  occurred,  while  in  1915  a  case  either 
slipped  through  undetected  or  was  a  visitor  infection; 
1916  two  cases,  1917  one  case,  1918  one  case  (see  orang- 
utan charts),  1919  and  1920  none  and  1921  one  case,  1922 
no  cases.  Since  1912  the  whole  monkey  collection  has  been 


THE  COMMUNICABLE  DISEASES 


539 


tested  every  two  years,  a  method  which  enabled  us  to  catch 
a  small  group  in  1916  and  has  protected  the  collection 
since  then.  Three  of  the  six  monkeys  specified  above  were 
never  placed  free  in  the  general  cages  of  the  exhibition 
house,  they  being  segregated  in  smaller  cages.   One,  the 


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Chart  P.    ORANG  UTAN  (Simia  satyrus).    Tuberculous. 

orang,  was  Avith  its  mate  in  an  isolated  cage.  The  other 
two  were  in  larger  cages  and  their  history  suggests  visi- 
tor infections. 

We  have  never  underestimated  the  possibility  that 
an  occasional  very  early  case  might  evade  detection  by 
this  test  but  we  believe  the  history  just  outlined  warrants 
us  in  depending  upon  it  for  the  protection  of  the  exhibi- 
tion. By  the  tuberculin  test  we  have  detected  the  existence 
of  the  disease  in  twenty-three  per  cent,  of  specimens. 


540  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Every  condemned  specimen,  forty-one,  showing  tuber- 
culosis, gave  a  positive  test.  Fifteen  monkeys  condemned 
on  their  temperature  charts  failed  to  show  the  disease. 
Eight  per  cent,  of  the  tests  resulted  in  suspicious  charts, 
and  the  animals  finally  died  of  the  disease  in  quarantine. 


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Chart  Q.    VARIEGATED  CEBUS  (Cebus  variegatus).    Tuberculous. 

Fifty-six  tuberculous  monkeys  died  on  exhibition,  of 
which  thirty-one  were  original  there  and  twenty-five  their 
contacts.  Twelve  of  the  thirty-one  were  in  the  early 
stages  of  the  work,  thirteen  due  to  our  misadventure  of 
1910  and  the  remainder,  six,  scattered  over  nine  years. 

Another  interesting  experience  concerns  the  exliibi- 
tion  of  a  group  of  Ehesus  Macaques  in  an  open  ''  band 
stand  "  cage.  The  idea  arose  in  an  attempt  to  find  a 
separate  exhibition  space  for  some  good  specimens  that 


THE  COMMUNICABLE  DISEASES 


541 


gave  unsatisfactory  charts,  with  the  purpose  of  applying 
at  the  same  time  the  *'  open  air  "  treatment  if  tubercu- 
losis existed.  The  experiment  has  been  entirely  success- 
ful since  in  the  eleven  years  during  which  this  enclosure 
has  been  used  there  has  been  but  a  single  case  of  tubercu- 


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Chart  R.    SOOTY  MANGABEY  (Cercocebus  fuliginoaus).    Tuberculous. 

losis  among  twenty-six  monkeys.  Curiously  enough  this 
exception  gave  a  good  chart  and  we  suspect  it  was  a 
visitor  infection ;  no  secondary  case  arose  from  it.  The 
animals  housed  in  this  cage  keep  in  excellent  condition, 
their  coats  responding  to  our  severe  winter  by  increasing 
in  thickness  and  glossiness.  Frozen  toes,  fingers  and 
tails  are  sometimes  seen  but  these  monkeys  seem  just  as 
happy  as  the  others.  Breeding  is  active  and  the  young 
are  lusty  and  husk^^  Practically  the  only  deaths  are  due 


542   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

to  accident,  or  to  abuse  of  old  and  less  vigorous  members 
of  the  colony.  We  are  unable  to  give  comparative  exhibi- 
tion periods  and  death  rates  for  monkeys  in  the  large 
house  and  tliis  open  cage  because  some  specimens  have 
been  changed  from  one  to  the  other  but  it  is  certain  that 


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Chart  S.    VERVET  MONKEY  (Cercopithecus  lalandii).    Tuberculous. 

the  appearance  of  the  *'  band  stand  "  monkeys  is  better 
than  those  in  the  house  and  there  are  four  of  eleven  ani- 
mals in  the  former  which  have  been  there  eleven  years 
and  only  four  among  the  seventy  in  the  exhibition  house 
for  that  length  of  time. 

The  results  of  the  foregoing  work  seem  to  demonstrate 
that  the  tuberculin  test  permits  the  separation  of  tubercu- 
lous and  non-tuberculous  monkeys  and  that  its  employ- 
ment   ser^^es    the    purpose    of    maintaining    a    healthy 


THE  COMMUNICABLE  DISEASES  543 

exhibition  by  excluding  infected  specimens.  These  experi- 
ences form  further  corroboration  of  the  facts  that  tuber- 
culosis begets  tuberculosis,  that  a  healthy  individual  is  not 
a  source  of  infection.  It  follows  that  an  obviously  tuber- 
culous animal  should  not,  need  not,  be  a  source  of  danger ; 


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Chart  T.    GRIVET  MONKEY  (Cereopithecus  sabaeua).    Tuberculoue. 

the  hidden  or  unrecognized  case  is  the  menace.  There 
is  little  or  no  problem  when  an  unequivocally  good  or 
bad  temperature  record  is  obtained ;  it  is  when  there  are 
slight  variations  from  the  standard  for  the  group  that 
decision  as  to  the  disposition  of  the  specimen  must  be 
made.  Nearly  always  such  specimens  are  retested  until 
the  records  are  definite.  If  they  be  constantly  irregular 
the  animal  is  either  sacrificed  or  exhibited  in  a  separate 
cage  far  from  other  monkeys.    It  is  by  the  sacrificing  of 


544  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

infected  specimens  or  the  segregation  of  suspected  ones 
that  our  collection  is  kept  clear  of  disease. 

Hygiene  of  a  general  character  must  be  maintained 
also.  Our  quarantine  rooms  are  disinfected  by  f ormalde- 
hj^de  and  mechanical  cleansing  after  every  case  detected 


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(See  page  533;. 


Non-tuberculous. 


as  tuberculosis,  and  painted  every  two  years.  Monkeys 
associated  with  infected  ones,  are  retested  and  then  given 
a  bath  of  carbolized  water  before  being  put  on  exhibition. 
The  exhibition  house  is  mechanically  and  chemically  dis- 
infected at  the  injection  time  each  two  years.  All  keepers 
are  examined  for  tuberculosis  upon  beginning  their  em- 
ployment and  those  handling  monkeys,  periodically  there- 
after.   When  a  case  of  tuberculosis  dies,  all  animals  in 


THE  COMMUNICABLE  DISEASES 


545 


the  same  and  adjoining  cages  are  removed  for  retest 
and  the  enclosure  scrubbed  and  disinfected. 

There  is  a  source  of  tuberculosis  upon  the  importance 
of  which  we  can  only  speculate — the  visitor.  There  were 
two  isolated  cases  in  animals  which  had  passed  the  test 


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Chart  V.    WEEPER  CEBUS  (Cebus  capucinus).    Tuberculous. 

with  unexceptionable  charts,  three  and  four  months  on  ex- 
hibition ;  curiously  enough  no  other  cases  occurred  in  their 
cages.  These  we  have  laid  to  visitor  infection  since  no 
previous  exposure  can  be  traced  for  the  specimens  and 
no  secondary  cases  occurred. 

I  can  conclude  this  discussion  of  the  tuberculin  test 
and  of  the  control  of  tuberculosis  by  its  use,  by  mention- 
ing the  possibilities  for  the  solution  of  the  problem  in  man, 
Wliile  the  eradication  of  the  disease  cannot  be  accom- 


546   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

plished  as  easily  as  if  a  potential  source  could  be  elimi- 
nated by  sacrifice,  it  will  come  in  direct  relation  to  the 
earliness  of  detection  of  infection  and  isolation  of  the 
sources  of  danger.  Not  so  much  the  cough-racked  con- 
sumptive but  the  unrecognized  early  lesion  whose  bearer 


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Chart  W.    CHACMA  BABOON  (Papio  porcarius).    Tuberculous. 

hawks  and  spits  in  public  places  or  at  home,  unaware  of 
his  malign  power! 

The  Skin  and  Eye  Tests  with  Tubeeculin. 

The  first  of  these  can  be  dismissed  briefly,  for  in  a  few 
cases  it  was  absolutely  of  no  value.  A  kno^ai  tuberculous 
monkey  was  injected  into  the  skin  of  the  chest  mth  0.5 
mg.  of  old  tuberculin.  The  small  bleb  disappeared  in  a  few 
hours  and  was  followed  by  no  reaction  whatsoever.  Other 


THE  COMMUNICABLE  DISEASES 


547 


attempts  likewise  failed,  some  of  them  I  believe  due  to  the 
technical  difficulty  of  injecting  into  the  skin.  This  tissue 
is  very  thin,  delicate  and  loose  at  the  less  hairy  places 
where  a  reaction  might  be  read — arm,  chest,  abdomen. 
The  hairless  parts  of  the  rump  might  be  used,  but  are  so 


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Chart  X.    BLACK  APE  (Cynopithecua  niger).    Tuberculous. 

often  scratched  and  soiled  with  dirt  that  readings  might 
be  misleading.  The  Von  Pirquet  test  was  done  on  the 
first  mentioned  specimen  and  was  likewise  negative.  His 
tuberculin  test  was  afterward  positive. 

The  ophthalmic  reaction  is  highly  spoken  of  in  the 
New  York  Zoological  Park  and  has  been  used  elsewhere. 
It  was  tried  by  me  at  the  time  Doctor  Blair  first  discussed 
it,  but  with  variable  results.  One  set  of  two  monkeys 
was  treated  with  Calmette's  purified  tuberculin  into  the 


548  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

conjunctival  sac  and  given  a  subcutaneous  dose  of  old 
tuberculin.  Another  set  received  1  per  cent,  old  tuberculin 
into  the  conjunctivae  and  the  usual  subcutaneous  dose. 
Although  all  these  monkeys  gave  a  temperature  reaction 
only  one  gave  a  conjunctival  reaction.  Fearing  that  the 
two  tests  simultaneously  might  be  an  unfair  trial,  another 
poor  specimen  was  given  an  eye  test  which  resulted 
negatively;  a  later  subcutaneous  test  and  autopsy 
revealed  the  disease.  Because  of  these  experiences  and 
the  fear  that  any  reacting  conjunctivae  might  become  sec- 
ondarily infected  from  the  uncertain  personal  hygiene  of 
the  beast,  we  decided  to  omit  this  method  and  rely  upon 
the  temperature  test. 

Pathological  Effects  of  the  Injection  of  Tuberculin'. 

It  is  generally  believed  that  tuberculin  injected  into 
tuberculous  animals,  in  doses  large  enough  to  produce  a 
marked  reaction  at  the  site  of  disease,  may  stimulate  the 
process  to  growth  and  spread  and  that  certain  parenchym- 
atous organs  in  such  bodies  undergo  degenerative 
changes.  We  can  give  little  information  concerning  the 
first  point  because  known  infected  animals  have  not  been 
sacrificed  during  the  test  and  we  are  not  informed  of  the 
degree  of  morbid  lesion  in  those  dying,  since  we  had  no 
previous  knowledge  of  its  existence. 

An  interesting  and  practically  useful  observation  has, 
however,  been  made  upon  the  kidneys  of  several  monkeys 
dying  shortly  after  tuberculin  injection.  It  consists  in  a 
marked  cloudy  swelling  of  the  renal  epithelium  and  a  con- 
gestion or  even  thrombosis  of  the  glomerular  capillaries, 
accompanied  sometimes  by  increase  of  nuclei  in  the  tuft 
and  by  amorphous  material  in  the  space  of  Bo^\^nan. 
Grossly  such  kidneys  are  but  little  changed,  albeit  the 
cortical  zone  may  be  dull  and  opaque  and  swell  out 
slightly  on  section;  very  occasionally  bloody  streaks  may 
separate  the  cortical  and  medullary  striae.    In  a  few  kid- 


Fl(,.  58.— EKFKCT  OF  TUBERCULIN  ON  THE  KIDNEY.  AN  UNUSUALLY  SEVERE  REACTION  IN  THE 
RENAL  TUBULES  AND  INTERSTITIAL  TISSUE  FOLLOWING  A  TEST  DOSE  OF  TUBERCULIN.  SUCH  A 
MARKED  CHANCE  SUGGESTS  THAT  IN  THIS  CASE  PRE-EXISTING  RENAL  DAMAGE  WAS  AGGRAVATED. 
THE   ANIMAL  WAS  TUBERCULOUS   BUT  HAD   NO   LESIONS   IN  THE  KIDNEY. 


THE  COMMUNICABLE  DISEASES  549 

neys  there  have  been  suggestions  of  preexisting  nephritis 
but  usually  the  findings  are  confined  to  those  given  above. 
At  all  events  true  glomerulonephritis  is  not  often  found. 
Monkeys  which  have  this  condition  may  or  may  not 
exhibit  a  behavior  suggesting  its  existence.  Sometimes 
it  will  be  noted  that  the  animal  is  dull  and  eats  little,  at 
other  times  the  keeper  mil  report  that  the  cage  is  seldom 
wetted  and  we  know  of  cases  in  which  only  an  ounce  or 
two  of  urine  has  been  passed  in  a  day.  Two  monkeys  were 
distinctly  ataxic  and  incoordinate  and  one  of  these  had  a 
convulsion.  From  one  a  specimen  of  urine  showed 
albumen  but  no  casts. 

These  signs  of  renal  affection  are  not  always  alone  nor 
are  the  kidneys  necessarily  the  only  part  diseased  since 
postmortem  records  show  a  variety  of  accompanying 
lesions,  bronchitis  and  enteritis,  for  example.  There  are, 
however,  several  cases  dying  in  a  few  days  after  tuber- 
culin injection,  both  with  and  without  tuberculosis,  in 
which  the  renal  changes  were  quite  prominent;  two 
examples,  without  tuberculosis,  exhibited  the  damage  to 
the  kidneys  very  well  and  with  no  other  evident  visceral 
pathology.  The  relation  of  cause  and  effect  may  not  be 
unequivocal,  but  these  findings  suggest  that  the  condition 
of  the  kidneys  deserves  attention  when  tuberculin  is  to 
be  injected.  My  associate,  Dr.  Corson-White,  is  firmly 
convinced  that  the  substance  whips  up  a  preexistent 
parenchjTuatous  disease  and  wants  to  see  a  urinalysis 
from  every  monkey  that  is  in  any  way  abnormal. 

The  Tuberculin-  Test  in  Other  Animals. 

Two  cases  of  tuberculosis  occurred  in  White-nosed 
Coatis  {Nasua  narica)  so  that  it  was  decided  to  test  their 
neighbors  in  the  next  cage.  There  is  reproduced  a  com- 
posite (Y)  of  the  temperature  record  of  three  of  these 
animals  after  receiving  2  mg.  of  tuberculin  under  the  skin. 
No  tuberculosis  was  found  in  them  at  death,  all  dying 


550  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

mthin  two  years.  The  similarity  to  the  primate  type  of 
temperature  curve  is  striking. 

Chart  Z  shows  the  course  of  temperature  before  and 
after  tuberculin  injection  in  a  Bactrian  Camel  (Camelus 
hactrianus)  2  which  lived  for  some  months  and  showed 
no  infection  at  autopsy. 

Charts  AA  and  BB  show  the  temperature  ranges  of 
respectively  a  healthy  and  a  tuberculous  Bison  [Bison 
bison.) 

Chart  CC  is  that  of  a  Malayan  Sambur  Deer  (Cervus 
equinus)  which  died  a  week  after  injection,  showing  fibro- 
caseous  tuberculosis.  The  failure  to  make  an  initial  rise 
is  noteworthy,  but  the  fall  in  temperature  may  be 
explained  by  the  severity  of  the  lesions  and  the 
approaching  death. 

Chart  DD  represents  daily  and  post-injection  records 
of  a  Virginia  Deer  (Cervus  virginianus)  which  at  death 
was  found  free  of  the  disease.  Cliart  EE  is  that  of  a 
healthy  American  Elk  {Cervus  canadensis) . 


THE  COMMUNICABLE  DISEASES  551 


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552   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 


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Chart  BB.    AMERICAN  BISON  (Bison  bison).    Tuberculous. 


THE  COMMUNICABLE  DISEASES 


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Chart  CC.     SAMBUR  DEER  (Cervus  equinus).     Tubercuk 


556  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 


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THE  COMMUNICABLE  DISEASES  557 


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Chart  EE.     AMERICAN  ELK  (Cervus  canadensis).     Non-tuberculous. 


SECTION  XVII— PART  2 
MYCOSIS 

This  is  a  general  term  applied  to  the  infections  with 
Hyphomycetes,  but  in  the  zoological  material  discussed 
here  it  refers  chiefly  to  the  growth  of  aspergillus  in  the 
air  sacs  and  viscera  of  birds. ;  A  few  cases  of  cutaneous 
mould  growth  have  been  seen  in  mammals  but  are  of 
trifling  importance ;  brief  mention  will  be  made  of  them 
on  a  later  page. 

Under  the  names  of  Aspergillosis  and  brooder  pneu- 
monia, the  infection  with  Aspergillus  fumigatus,  A.  glau- 
cus  and  others  is  well  known  to  breeders  of  chickens, 
ducks  and  ostriches.  Literary  references  to  its  occurrence 
in  zoological  collections  are  numerous,  but  there  seems  to 
be  no  record  indicative  of  its  frequency  in  the  various 
orders  nor  discussion  of  the  pathological  types  best 
exhibited  by  different  birds.  There  has  indeed  been 
some  discussion  of  the  actual  entity,  mould  disease,  the 
picture  found  at  autopsy  being  referred  to  secondary  con- 
tamination with  fungi  in  the  presence  of  bacterial  infec- 
tion. Experiments  by  DeLong  and  others  have  been 
indifferently  successful  in  the  production  of  the  disease 
by  iiilialation  of  mould  spores.  The  appearance  of  groups 
of  cases  in  breeding  places,  apparently  all  exhibiting  the 
same  organism,  in  the  absence  of  other  varieties  of  avian 
epizootics  seems  to  warrant  the  conclusion  that  the 
hyphomycetes  can  at  least  be  associated  with  a  fatal 
morbid  lesion  of  quite  uniform  character  whether  or  not 
they  be  the  original  invaders. 

Judging  by  our  experience  it  would  seem  probable  that 
the  aspergillus  can,  under  conditions  not  fully  explained, 
cause  inflammation  of  the  avian  air  sac  and  tubercles  in 
viscera,  in  the  absence  of  other  evident  causes  of  illness 
and  death.    Over  three-fourths  of  our  cases  have  no  other 

558 


MYCOSIS  559 

diagnosis  than  ''mould  disease";  this  may  be  in  part 
due  to  overlooking  other  things,  but  to  a  much  greater 
extent  to  the  very  extensive  mould  growth  which  obscures 
all  other  changes.  That  unexplained  preparatory  condi- 
tions may  exist  is  indicated  by  general  observation  and 
some  experimental  work.  Whereas  in  breeding  estab- 
lishments this  disease  occurs  in  epizootics,  or  in  groups 
of  cases,  with  us  it  is  enzootic,  constantly  present,  never, 
however,  bursting  forth  in  virulent  form  with  high 
mortahty.    Nor  is  the  condition  highly  contagious. 

In  an  attempt  to  explain  its  source  I  examined  over 
forty  varieties  of  feed  and  found  therein  several  strains 
of  aspergillus  and  of  mucor ;  the  latter  occurs  occasionally 
in  the  avian  air  sac.  The  infective  material  is  therefore 
constantly  present,  and  it  would  seem  that  if  it  were 
capable  of  initiating  a  fatal  disease  many  more  cases 
should  come  to  our  attention.  Perhaps  these  moulds  do 
gain  access  to  the  avian  air  sac  and  are  killed  off,  or  only 
assume  a  pathogenic  role  when  they  are  in  large  num- 
bers or  a  preexisting  disease  assists  them.  Since  our 
records  do  not  support  the  idea  that  a  preceding  condi- 
tion must  exist  for  a  growth  of  mould  to  be  successful, 
and  yet  pathological  and  experimental  observations  sug- 
gest that  something  helps  its  colonization  in  the  air  sac, 
what  are  such  conditions!  Moulds  grow  on  feed  and  lit- 
ter in  which  birds  pick ;  from  this  it  is  quite  possible  for 
a  piece  of  grain  or  even  inorganic  matter  laden  with 
spores  to  be  inhaled  and  lodged  in  a  secondary  alveolus 
near  the  air  sac  whence  extension  into  the  air  spaces  could 
occur.  In  addition  I  think  it  quite  conceivable  that  a 
whole  colony  of  mould  might  be  inspired  with  the  same 
result,  the  mechanical  obstruction  being  sufficient  physical 
damage  to  incite  inflammation.  The  continuous  moist 
surfaces  of  the  bronchial  passages  and  air  sacs  afford 
conditions  favorable  to  the  growth  of  mould  and  as 
inflammation  is  not  vigorous,  little  resistance  is  presented 
to  its  spread. 


560  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

The  mode  of  operation  of  these  hyphomycetes  has 
usually  been  assumed  to  be  a  mechanical  one,  local 
colonization  replacing  healthy  tissue  or  spreading  along 
surfaces  so  that  function  is  physically  impossible.  A 
support  of  this  idea  is  to  be  found  in  the  fact  that  inflam- 
mation, as  produced  by  schizomyces,  is  trifling  or  absent; 
the  necrosis  that  occurs  is  due  to  choking  off  of  tissue  by 
the  intricately  tangled  masses  of  mycelia  and  blocking 
off  of  air  or  blood  supply.  The  existence  of  an  infiltrating 
and  necrotizing  form  in  some  parrots  and  gallinaceous 
birds,  suggested  to  me  that  a  toxin  might  be  responsible 
for  some  part  of  mould  action.  Proof  for  this  speculation 
was  sought  by  injecting  into  the  pectoral  muscles  of 
pigeons  an  emulsion  of  a  dead  mould  and  a  filtered 
broth  culture.  Necroses  occurred  but  only  to  an  extent 
which  I  interpreted  as  due  to  the  physical  destruction  of 
muscle  by  the  injected  material ;  they  were  larger  with  the 
dead  mould  than  with  broth  filtrate.  I  concluded  there- 
fore that  aspergillus  perhaps  has  no  toxin  as  usually 
described  for  bacteria. 

Types  of  Mycosis. 

Avian  mycosis  occurs  in  three  different  forms,  two  of 
which  are  probably  of  similar  nature  and  two  are  fre- 
quently combined.  (The  first  variety,  most  often  seen  in 
gallinaceous  and  anserine  birds,  consists  of  thickening 
and  opacity  of  the  air  sac  walls,  upon  the  surface  of  which 
either  a  curd-like  pseudocoagulum  or  a  velvety  or  fluffy 
mould  growth  appears.  This  variety  usually  begins 
about  the  anteroinferior  pulmonary  stoma  on  the  right 
side  extending  thence  to  the  related  sac,  upward  toward 
the  wing  and  downward  to  the  abdominal  spaces. 
Occasionally  the  middle  thoracocervical  space  is  involved, 
probably  via  the  opening  in  the  syrinx.  Extension  takes 
place  by  the  way  of  normal  passages,  but  when  the  growth 
is  dense  it  also  seems  to  occur  by  continuity  through  tis- 
sue.   This  variety  may  or  may  not  be  associated  with  the 


Fig.  59. — NODULAR  OR  TUBERCULAR  MYCOSIS  IX  THE  LUNGS  OF  A  DUCK.  ONE-HALF  OF 
THE  LUNG  IS  CUT  AWAY  AND  LAID  ON  THE  INTESTINES  TO  CONTRAST  THE  PLEURAL  AND 
SECTION  SURFACES. 


MYCOSIS  561 

second,  an  infiltrative  type  of  lesion  best  seen  in  the  lung. 
Under  what  seems  to  be  a  true  picture  of  mycotic  pneu- 
monia, dirty  gray  consolidated  areas  will  be  found  around 
the  bronchial  space,  infiltrating  in  all  directions  and  with- 
out definite  boundaries.  A  similar  lesion  has  been  seen 
also  in  the  liver  on  rare  occasions,  but  the  lung  is  its  usual 
seat.  Judging  from  microscopic  appearances  this  is  a 
process  complicated  by  the  addition  of  bacteria. 

l^  The  third  variety  is  nodular  or  tubercular  mycosis,  a 
process  of  probably  more  chronic  nature  since  around  the 
isolated  lesions  connective  tissue  is  perceptible,  it  being 
absent  or  inconspicuous  in  the  other  types.  The 
formation  of  gray  or  yellow-white  nodules  from  a  few 
millimetres  to  a  centimetre  in  cross  section,  is  the  charac- 
teristic production  in  this  variety.  Lungs,  liver,  spleen, 
intestines  and  air  sacs  are  involved  in  about  this  order. 
Attempts  at  explanation  of  this  peculiarity  of  growth 
were  made  in  the  direction  of  identification  of  the  species 
of  mould,  kind  of  bird  and  probably  degree  of  resistance. 
No  conclusions  could  be  drawn  since  the  same  variety  of 
mould  was  found  in  this  as  in  other  types ;  no  bird  showed 
a  special  susceptibility  or  resistance  to  it.  It  is  quite 
diflBcult  to  obtain  a  culture  from  nodular  mycosis,  it  being 
necessary  to  crush  or  grind  the  solid  masses  before 
making  cultural  implants.  This  variety  should  always 
be  differentiated  from  tuberculosis  by  staining  for 
the  organisms,  j 

Histologically  studied  these  three  types  are  not  as 
easily  separated  as  the  gross  appearances  would  warrant 
one  to  expect.  The  original  mould  nodule  begins  in 
essentially  the  same  manner  in  all,  a  small  colonization 
of  mycelia  and  spores  which  grow  centrifugally,  but  ever 
becoming  more  intricately  wound  in  their  first  location. 
As  the  tissue  is  invaded,  total  necrosis  takes  place,  no 
recognizable  cellular  architecture  being  left.  When  fully 
formed  the  mycotic  tubercle  consists  from  the  centre 
outward  of  a  necrotic  mass,  in  which  spores  and  mycelia 


562   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

stain  indifferently  well,  surrounded  by  a  dense  zone  of 
fully  formed  mould  beyond  which  mononuclear  cells  and 
a  few  loose  fibrils  may  be  found.  The  circumferential 
tissue  of  the  viscus  supplies  a  mild  congestion  and  per- 
haps a  moderate  connective  tissue  capsule.  Giant  cells, 
as  known  for  tuberculosis,  do  not  appear. 

Where  the  process  involves  loose  tissue  like  the  lung, 
especially  when  growing  rapidly,  the  sporulating  heads 
of  the  mycelia  stretch  out  in  advance  of  the  main  mass 
and  resemble  rays.  So  too  in  a  rapidly  gromng  nodule 
radiating  mycelia  are  sometimes  seen  but  never  with  the 
regularity  of  arrangement  typical  for  actinomycosis. 
Diffuse  and  irregular  mycelial  spread  is  characteristic 
of  the  second  or  infiltrative  type  of  lesion  and  between  the 
mould  stalks  one  may  discover  well-preserved  tissue  cells 
and  at  times  bacterial  forms  like  cocci  or  bacilli.  This 
picture,  suggesting  as  it  does  bacterial  admixture  and 
more  active  inflammation,  leads  one  to  the  conclusion  that 
the  morbid  process  in  which  it  is  found  does  not  represent 
mould  disease  per  se,  but  a  mixed  infection.  Whether  or 
not  the  bacterial  disease  exists  first  and  paves  the  way  for 
the  mould  I  am  unprepared  to  say;  I  am  inclined  to  the 
view  that  mycosis  can  start  by  itself. 

iNcmENCE  IN  Avian  Orders. 

Mycosis  may  be  said  to  occur  in  all  birds  although  the 
appended  list  fails  to  show  cases  in  a  few  of  the  orders 
included  in  this  study;  the  sum  total  of  autopsies  from 
missing  orders  is  only  45,  so  that  they  may  be  ignored. 


Picarise. . . 
Psittaci .  . 
Striges.  .  . 
Accipitres. 
Columbae. 
Galli 


3.7 
1.2 
3.9 
6.7 
5.1 
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2.7 


Fulicarise .  . 
Alectorides. 

Gaviae 

Impennes.  . 
Herodiones . 
Anseres. . .  . 
Struthiones . 


8.6 

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10. 
40. 

2. 

6. 

9.4 


For  meaning  of  italics  see  foot  note  Table  i. 


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IG.   60. — MICROSCOPIC   APPEARANCE    OF   ONE    OF    THE    NODULES.      NOTE    THE    CENTRAL 
NECROSIS  AND  THE  CLOSE   FIBROCELLULAR   PACKING   IN   THE  RESTRAINING  CAPSULE. 


MYCOSIS  563 

These  figures  hardly  permit  conclusions  as  to  rela- 
tive vulnerability  unless  the  large  percentage  of  cases  for 
the  small  number  of  Impennes,  Gavise  and  Struthiones  be 
permitted  to  stand.  Judging  from  orders  upon  which 
there  are  at  least  one  hundred  autopsies,  owls,  ducks  and 
eagles  are  most  likely  to  suffer  with  mycosis.  The  per- 
centages are,  however,  not  very  convincing,  and  it  would 
seem  better  with  these  data  to  conclude  for  the  present 
that  any  variety  of  bird  is  susceptible  to  mould.  Ostriches 
have  long  had  the  reputation  of  succumbing  to  this  affec- 
tion, so  that  their  outstanding  position  in  the  list  is  more 
easily  credited. 

It  was  to  be  hoped  that  the  various  susceptibilities 
would  assist  in  an  explanation  of  the  genesis  of  mould 
disease,  but  the  result  of  the  analysis  is  suggestive  only 
in  one  direction  which  can  be  stated  quite  briefly. 
Ostriches,  owls,  shore  and  swimming  birds  have  large 
stomata  between  the  lungs  and  the  lateral  air  sacs  and 
show  a  high  mould  disease  incidence.  The  natural  thought 
is  that  access  of  mycelia  and  spores  to  the  air  sacs  is 
facilitated.  More  comparative  data  is  being  assembled 
upon  this  point.  Water  birds  are  more  susceptible  to 
mould  than  land  birds ;  the  percentages  based  upon  cases 
and  totals  for  orders  is  5.4  to  3.7. 

The  hygiene  of  mycosis  is  that  of  scrupulous  cleanli- 
ness. Being  hampered  by  incomplete  knowledge  as  to 
its  genesis  one  can  only  apply  common  sense  measures. 
The  germs  having  been  found  upon  all  the  vegetable 
feeds,  it  naturally  follows  that  they  cannot  be  eradicated, 
but  their  colonization  in  large  numbers  can  be  prevented 
by  repeated  cleansing  or  sterilization  of  bins  and  pans 
so  that  no  mouldy  or  musty  material  is  given  to  the 
animals.  At  times  of  serious  outbreaks  sterilization  by 
burning  all  old  feed,  starting  fresh  with  good  material  in 
bins  painted  or  saturated  with  disinfectant  and  then 
deodorized,  is  the  only  salvation.  Where  the  blast  lamp 
can  be  applied,  it  is  the  safest  procedure.     Autoclave 


564   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

sterilization  should  be  thorough  if  practiced.  B}'  con- 
stant vigilance  we  believe  that  our  large  exhibition  house 
is  protected,  but  the  sanitation  of  the  pond,  where  many 
ducks  have  the  disease,  can  never  be  so  satisfactory. 

Mycosis  among  mammals  as  an  organic  pathological 
entity  is  certainly  a  rare  condition,  indeed  almost  always 
to  be  considered  an  accidental  or  secondary  one.  In  so 
far  as  human  pulmonary  disease  is  concerned  it  is  among 
the  pathological  curiosities  deserving  of  individual 
report.  Moulds  of  several  varieties  have  been  found  in 
intestinal  ulcers,  in  cranial  sinuses  and  in  the  ear,  while 
generalized  mycosis  from  thrush  and  favus  are  reported 
(cases  of  thrush  in  kites  with  fatal  outcome  has  been 
described  on  page  168).  Cutaneous  infestation  with 
mould  is  quite  another  matter  for  ring  worms  and  similar 
conditions  are  now  believed  to  be  due  solely  to  the  pene- 
tration by  spores  and  mycelia  into  the  superficial  dermis 
and  into  hair  follicles.  Numerous  small  lesions  have  been 
detected  on  dogs  and  cats  that  coiTespond  to  the  ring 
worms  described  for  them ;  to  these  we  have  devoted  little 
study,  because  when  discovered  the  animal  is  removed, 
treated  or  killed  to  protect  others.  Doctor  Weidman  has 
discovered  several  varieties  of  hyphomycetes  which  he 
will  report  upon  at  a  later  time.  No  case  of  generalized 
mould  disease  has  been  found  in  mammals,  but  Doctor 
Weidman  has  discovered  a  hitherto  undescribed  mould 
in  ulcers  and  cutaneous  abscesses  of  seals;  no  ex- 
tension beyond  subcutaneous  areolar  tissue  occurred  in 
these  cases. 

Botryomycosis,  while  not  strictly  belonging  to  the 
foregoing  group,  may  be  mentioned  here  because  of  its 
nodular  tumor-forming  superficial  growths.  The  case  to 
be  cited  certainly  belongs  to  this  illy  defined  group  of 
diseases  even  if  the  organism  was  not  isolated.  It  cannot 
be  accepted  as  wholly  demonstrated  that  the  disease 
described  under  this  name  is  always  the  same  or  that  it 
has  a  single  cause. 


MYCOSIS  565 

California  Hair  Seal  {Zalophus  calif ornianus)  9  .  Had  sore  spots 
on  side  for  several  months.  Ate  well  up  to  three  days  before  death 
but  ate  something  to  time  of  death. 

Diagnosis. — Chronic  enteritis  Avith  acute  hemorrhagic  exacerbation. 
Low  grade  chronic  diffuse  nephritis.  Botryomycosis.  General  condi- 
tion poor,  subcutaneous  fat  practically  absent.  On  the  left  side  of  the 
thorax  there  is  a  warty  and  nodular  thickening  of  the  skin  overlying 
diffuse  and  flat  thickening  of  subcutaneous  tissues.  There  are  warty, 
pustular,  fistulous  communications  in  four  places  between  surface  and 
deep  mass.  They  are  covered  over  with  light  crust.  On  dissection 
mass  is  found  to  be  in  subcutaneous  tissue  well  outlined  and  encapsulated 
and  consisting  on  section  of  dense,  white,  firm  trabecul83  forming  a 
mesh  around  yellow,  soft  areas  which  can  be  squeezed  out.  An  adjoin- 
ing lymph  node  is  much  enlarged,  dense,  tough,  resilient,  on  section 
showing  great  connective  tissue  increase  and  solid  brown  medulla.  The 
thyroids  are  solid,  brown,  3.  x  2.  x  .6  cm.  and  3.7  x  l.S  x  .6  cm.  Trachea 
and  bronchi  contain  pink  froth  but  mucosa  is  negative.  Anterior  edges 
of  lungs  are  distinctly  emphysematous.  Remainder  of  lung  is  uni- 
formly congested  and  lobules  are  quite  prominent.  No  consolidations. 
Bronchial  lymph  nodes  are  slightly  large,  anthracotic  and  wet.  The 
heart  is  dilated,  filled  with  mixed  clot,  muscle  firm  and  deep  brown 
color.  The  liver  surface  is  smooth,  edges  sharp,  size  normal,  color 
dull  brown  mth  greenish  cast,  consistency  firm  and  tough.  Section 
surface  is  glistening,  smooth,  moist,  lobules  indistinct  but  probably 
normal.  The  gall-bladder  is  full  of  fluid  yellow  bile.  The  common 
duct  is  patulous.  The  spleen  is  of  normal  size  and  shape,  capsule  opaque 
gray.  The  trabeculae  are  prominent,  the  pulp  stippled  rusty  brown. 
The  kidney  is  of  normal  size,  capsule  is  smooth,  strips  easily  leaving  a 
smooth  purple  surface.  The  organ  is  firm.  The  lobules  and  lobular 
markings  are  distinct.  The  adrenal  has  a  narrow  regular  brown  cortex 
and  gray  homogeneous  medulla.  The  stomach  contains  whole  fish. 
The  mucosa  shows  digestion,  congestion  and  mucus  formation.  Begin- 
ning at  the  pyloinis  and  extending  to  the  colon  the  mucosa  is  swollen 
and  edematous,  yellow  brown.  In  the  lower  part  it  is  quite  firm  and 
opaque.  In  the  upper  part  it  is  more  translucent  except  where  there 
are  difl'use  hemorrhagic  mottlings  of  the  submucosa.  Here  and  there 
are  shallow  erosions  but  no  ulcers.  The  lymphatics  of  tlie  mesentery 
are  definitely  enlarged,  white,  pale  and  very  firm.  Smear  from  the 
surface  of  the  growth  in  side  fails  to  show  any  definite  yeasts  or  moulds 
by  Loeffler's  or  Gram's  stains.  It  is  largely  made  up  of  polynuelear 
cells  mth  many  large  mononuclears,  many  of  which  are  pha^oeyting 
polynuclears  and  nondescript  bodies.  Cultures  failed  to  grow.  Micro- 
scopic section  of  liver  shows  marked  congestion  with  slight  hydropic 
degeneration  of  the  epithelium.  The  kidney  capsule  is  not  thickened. 
Interstitial  tissue  not  grossly  exaggerated  but  connective  tissue  nuclei 
fairly  numerous.     Some  tufts  have  decidedly  more  enlongated  nuclei 


566   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

than  others  and  connective  tissue  around  the  stalk  vessels  seems  hyaline. 
Capsule  for  mcjst  part  not  thickened  but  space  contains  cells  and 
detritus  in  many  instances.  Epithelium  of  the  tubules  for  the  most 
part  swollen,  loosened  and  without  nuclei.  Some  imperfectly  formed 
casts.  The  intestine  shows  distinct  congestion  of  the  whole  mucosa 
with  here  and  there  definite  small  hemorrhages  well  out  in  the  villi. 
Slight  round  cell  increase  but  no  definite  fibrosis.  In  submucosa  aroun<l 
vessels  connective  tissue  is  hyaline  in  many  places.  The  tumor  on  side 
consists  of  dense  strands  of  connective  tissue  forming  alveoli  of  vary- 
ing size  containing  an  exudate  of  fibrin  and  cells,  about  three-fourths 
of  the  latter  being  mononuclears.  Neither  connective  tissue  nor  cells 
are  arranged  in  a  characteristic  or  peculiar  manner  so  the  observer 
is  forced  to  conclude  that  this  is  one  of  the  conditions  of  the  group 
called  Botryomycosis. 


SECTION  XVII— PART  3 
THE  STREPTOTHRICOSES 

The  organisms  belonging  to  the  genera  Streptothrix, 
Actinomyces,  Discomyces  and  Nocardia  as  named  by 
various  authors  have  in  common  the  power  to  produce 
local  chronic  inflammation  of  gradually  spreading  char- 
acter and  chronic  course.  Their  most  conspicuous  repre- 
sentative, the  ray  fungus,  is  best  known  as  the  producer 
of  lumpy  jaw  in  cattle  and  as  an  occasional  pathogen  in 
man.  Other  members  of  the  group  cause  certain  lymph- 
channel  disease  in  domestic  animals  and  pulmonary  dis- 
ease in  man.  Pathogenic  power,  it  is  believed,  lies  in 
the  ability  of  these  organisms  to  colonize  and  irritate, 
thus  producing  continuously  enlarging  tumefactions,  no 
evidence  being  at  hand  that  any  of  them  produce  a  toxin 
either  in  their  surroundings  or  within  their  own  bodies. 
Because  of  their  constant  irritation,  bacterial  mixed  in- 
fection often  ensues  so  that  purulent  degeneration  may 
occur  at  the  original  site  of  disease  and  thence  may  spread 
via  the  blood  vessels,  or  by  continuity  of  tissues  or,  if  the 
mucous  membrane  of  the  pharynx  be  diseased,  by  the  air 
passages. 

The  study  of  the  genesis  of  actinomycosis  is  by  no 
means  a  closed  one.  While  it  is  believed  that  pastures 
and  fodder  carry  the  organism  and  that  it  gains  access 
to  the  tissues  by  passing  into  small  wounds  that  are  made 
by  sharp  sticks  or  grain  beards,  the  exact  origin  of  the 
disease  is  not  understood.  The  original  lesion  is  cer- 
tainly trifling  and  the  fully  developed  one  may  not  be 
discoverable  until  it  is  well  under  way  and  causes  external 
deformity.  Even  when  sloughing  has  occurred,  the  dis- 
ease is  not  very  communicable.  The  method  of  contrac- 
tion of  lymphatic  streptothricosis  in  cattle  is  believed  to 
be  from  other  cases  via  skin  wounds  or  if  abrasions  be 

567 


568   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

soiled  by  infective  dirt.  Just  how  human  beings  contract 
these  infections,  in  the  absence  of  infected  cattle  is  un- 
known, but  for  the  pulmonary  form  the  route  usually 
followed  in  tuberculosis  is  probably  taken. 

The  material  of  our  zoological  collection  permits  few 
observations  of  value  upon  "  lumpy  jaw  "  but  we  have 
encountered  a  streptothricosis  of  kangaroos  which  may 
throw  some  light  upon  the  whole  subject  and  to  these  cases 
I  shall  devote  considerable  space  since  no  description  of 
it  occurs  in  the  literature. 

Actinomycosis. 

This  disease  has  been  diagnosed  with  certainty  in  two 
American  Tapirs  and  with  reasonable  satisfaction  in 
three  deer.  Two  Malayan  Tapirs  have  also  had  lumpy 
jaw  clinically  but  the  organisms  were  not  found.  It  is  in- 
teresting and  noteworthy  that  other  zoological  collections 
have  observed  the  disease  in  this  same  animal,  a  fact 
which  suggests  the  high  susceptibility  of  the  tapir  to 
actinomycosis.  There  are  recorded  in  the  protocols 
a  few  times  sluggish  ulcers  on  the  tongue  in  other  ungu- 
lates but  I  am  not  prepared  to  label  them  as  actinomycotic 
since  on  one  occasion  smears  and  sections  were  studied 
with  great  care  and  nothing  found  to  justify  such  a  diag- 
nosis; nor  were  there  maxillary  or  pulmonary  lesions. 
Before  passing  to  a  discussion  of  the  diagnosis  and  mor- 
bid characters  it  seems  worthy  of  emphasis  that  our  cases 
of  this  disease  should  appear  in  one  family  of  Perissodac- 
tyla  and  in  one  family  of  Artiodactyla,  in  the  latter 
not  affecting  Bovidae,  the  family  to  which  domestic 
cattle  belong. 

The  diagnosis  of  lumpy  jaw  depends  upon  the  growth 
of  tumors  in  the  neck  and  maxillary  regions  wliich  tend 
to  break  down  and  discharge  a  thick  pus  containing 
** sulphur  granules,"  little  masses  of  necrotic  matter  sur- 
rounding colonies  of  the  ray  fungus.  When  these  condi- 
tions are  fulfilled,  the  matter  is  easily  enough  settled. 


THE  STREPTOTHRICOSES  569 

This  was  possible  with  the  tapirs  but  in  the  deer  the  con- 
clusion was  not  so  easily  reached  and  the  diagnosis  had 
to  be  made  partly  by  exclusion.  Anatomically  the  gross 
and  microscopic  appearances  of  lumpy  jaw  in  the  tapirs 
follows  the  text-book  descriptions  but  our  cases  in  the 
deer  deserve  separate  comment.  Whether  or  not  these 
differences  mean  a  peculiarity  of  resistance  on  the  part 
of  the  animal  or  a  new  variety  of  streptothrix  only  fur- 
ther study  can  settle. 

The  beginning  of  the  lesion  in  the  deer  was  in  the  jaw 
bone  as  circumscribed  or  fusiform  swellings  appearing  on 
the  under  surface.  Growth  usually  progressed  into  the 
pharyngeal  cavity  and  backward  under  the  ear,  but  a  large 
tumor  stretcliing  down  the  neck  was  only  observed  once. 
Suppuration  and  ulceration  occurred  tmce  but  only  once 
were  bacteriological  observations  possible  before  death, 
and  then,  they  were  negative.  The  fatal  outcome  seemed 
to  be  due  to  inanition,  possibly  because  the  animal  could 
not  eat,  for  respiratory  tract  involvement  was  only  pres- 
ent once  and  then  to  a  trifling  degree.  At  autopsy,  actino- 
myces  in  ray  form  were  found  in  one  animal  only,  the 
diagnosis  resting  upon  histology  in  the  other  two.  Nor 
did  the  degenerated  centre  of  the  swelling  contain  the  sul- 
phur granules  in  any  case. 

The  microscopic  characters  of  the  tumors  resembled 
those  of  giant  cell  sarcoma  and  chronic  rarefying  osteo- 
periosteitis  mth  areas  of  round  cell  infiltration  but  no 
granulomata  as  are  occasionally  seen  in  lumpy  jaw.  I 
have  always  felt  that  a  ''  giant  cell  sarcoma  "  with  in- 
flammation when  seated  in  the  jaw  of  a  lower  animal 
should  be  looked  upon  with  grave  suspicion  and  be 
searched  diligently  for  fungi.  The  organisms  could  not 
be  found  in  sections  of  any  of  these  cases,  although  pres- 
ent in  the  pus  from  a  pocket  in  one.  Reference  has  been 
made  in  discussing  tumors  of  the  bones  in  gazelles  and 
opossums  to  their  resemblances  to  osteofibroma  and  ac- 
tinomycosis. The  diagnoses  were  made  after  long  study 
37 


570  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

of  the  notes  and  sections.  Actinomycosis  is  usually  uni- 
lateral while  leontiasis  ossium  is  commonly  bilateral; 
the  tumors  have  not  broken  down  nor  spread  into  the 
neck.  In  one  macerated  jaw  bone  the  osteoporosis  and 
hypertrophic  periosteitis  were  comparable  to  those  of  the 
bo\ane  form  but  the  masses  were  not  so  extensive  as 
is  common  for  domestic  cattle. 

Treatment  of  this  disease  was  attempted  in  the  tapirs 
but  not  in  the  deer;  the  latter  are  too  nervous  to  be 
handled  repeatedly  with  safety  to  themselves.  Follow- 
ing the  usual  method,  potassium  iodide  was  administered 
in  saturated  solution  on  bread,  beginning  at  twenty  grains 
thrice  daily  and  rising  in  twoi  cases  to  sixty  grains  thrice 
daily.  It  cannot  be  stated  that  any  material  improve- 
ment followed  this  heavy  dosage  although  in  one  case 
the  disease  was  very  protracted — some  six  months,  so 
that  it  may  have  modified  the  progress  of  the  lesion. 
However,  other  things  were  done  for  the  beast  so  that 
the  effect  of  any  one  kind  of  treatment  is  difficult  to 
evaluate.  It  was  noted  that  iodide  served  to  keep  the 
stools  quite  loose  and  that  its  withdrawal  was  followed 
by  constipation;  upon  resumption  of  the  drug  normal 
bowel  movements  appeared.  Every  soft  spot  was  opened 
surgically  to  allow  the  pus  to  drain  away.  A  vaccine  of 
Act.  bovis  was  prepared  and  injected  under  the  hide  be- 
ginning at  0.5  mg.  and  rising  to  2.5  mg.  in  five  doses  after 
which  the  animal  became  so  unruly  that  the  injections  had 
to  be  discontinued.  On  the  whole  we  are  not  impressed 
with  the  probability  of  success  in  the  treatment  of  actino- 
mycosis in  tapirs.  In  the  future  we  propose  to  try  opera- 
tion and  the  use  of  Dakins  solution  or  Dichloramin  T. 

Streptothricosis  or  Nocardiosis  or  Kangaroos. 

A  fatal  disease  of  Australian  marsupials  character- 
ized by  swellings  and  ulcerations  about  the  lips,  teeth, 
tongue  and  cervical  tissues  is  known  apparently  all  over 
the  world  by  observers  of  these  animals  in  collections. 


THE  STREPTOTHRICOSES  571 

From  commercial  shippers  of  animals,  from  zoologists 
and  naturalists  we  have  reports  that  wherever  kangaroos 
and  wallabies  are  exhibited  this  disease  makes  its  ap- 
pearance and  carries  off  a  considerable  percentage  of 
the  collection.  A  fully  developed  case  bears  a  noteworthy- 
resemblance  to  lumpy  jaw,  being  called  "  jaw  disease  " 
by  non-medical  observers.  However,  it  is  highly  probable 
that,  while  the  most  conspicuous  morbid  changes  occur 
around  the  jaw,  the  agent  provocative  of  the  disease  is 
capable  of  causing  different  pathological  effects  and  that 
certain  cases  of  septicemia  and  gastroenteritis  are  due 
to  it ;  Doctor  Blair  of  New  York  concurs  in  this  opinion. 
Our  study  of  the  problem  would  inculpate  a  variety  of 
Nocardia,  possibly  assisted  by  certain  schizomyces.  I 
have  seen  in  the  literature,  but  unfortunately  cannot  lo- 
cate, a  reference  to  an  article  by  a  Russian  who  observed 
the  disease  and  was  con\iLnced  that  its  cause  is  to  be  found 
in  a  streptothrix(l)  obtainable  from  the  necroses  in  the 
soft  tissues  of  the  jaw,  a  view  entirely  in  accord  with 
our  findings. 

The  disease  is  not  very  communicable  because  its 
appearance  in  a  pen  need  not  be  followed  by  secondaries 
in  the  mates  of  the  sick  beast.  It  appears  chiefly  in  newly 
acquired  specimens  but  may  develop  sporadically  in  those 
exhibited  a  long  time  and  apparently  not  associated  with 
recent  acquisitions.  This  suggests  two  or  three  possi- 
bilities. It  may  be  imported  by  new  arrivals,  or  new- 
comers may  meet  a  germ  to  which  they  are  unaccustomed 
and  therefore  less  resistant,  the  strange  surroundings 
reducing  their  opposition  to  it.  Old  specimens  may  have 
enough  resistance  to  withstand  infection  entirely  or  only 
succumb  to  large  doses.  Lastly  one  comes  to  the  explana- 
tion commonly  employed  for  actinomycosis,  the  presence 
of  the  organisms  in  fodder  or  pasture,  perhaps  all  the 

( 1 )  This  term  will  be  used  in  the  following  pages  to  mention  the 
organism  since  by  many  persons  it  is  better  known  than  Nocardia  and 
moreover  describes  the  form  better.  I  believe  genus  Nocardia  is  the  correct 
nomenclature  for  reasons  given  on  a  subsequent  page. 


572  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

time,  but  gaining  entrance  to  the  animal's  body  via 
wounds  made  by  sharp  sticks  or  the  beards  of  grain. 

While  circumstantial  evidence  offers  some  support  to 
this  general  idea,  it  cannot  be  accepted  as  proven.  I 
have  not  made  studies  of  the  feed  for  the  purpose  of 
isolation  of  the  streptothrix  but  cultures  from  the  lips  and 
gingival  margin  of  healthy  and  infected  animals  were 
made  for  its  cultivation.  These  attempts  were  fruitless, 
and  without  wonder  since  the  germ  when  isolated  from 
a  fully  developed  case  is  quite  finical  in  its  manner  of 
growth;  there  are  so  many  kinds  of  bacterial  life  that 
they  may  easily  overgrow  the  one  in  quest.  So,  too,  cul- 
tures made  directly  from  subcutaneous  necrotic  areas  may 
not  always  give  a  positive  growth  although  smears  from 
the  same  material  may  reveal  numerous  threads  under 
the  microscope. 

The  idea  that  sharp  grasses  are  responsible  for  the 
origin  of  Kangaroo  disease  finds  a  protagonist  in  Dr.  A. 
S.  LeSouef,  Director  of  the  Zoological  Garden  at  Sidney, 
Australia,  a  gentleman  whose  judgment  carries  weight. 
He  writes  :  ''We  have  found  that  it  is  entirely  due  to  get- 
ting spear  or  barley  grass  in  their  food;  omng  to  the 
formation  of  the  mouth,  this  grass  gets  wedged  in  be- 
tween the  teeth  and  the  cheeks,  penetrates  the  flesh  and 
allows  the  bacteria  to  get  a  footing,  this  in  time  heaps  up 
on  the  inside  and  forms  an  abscess  that  bursts  exteriorly. 
Formerly  all  the  Australian  Zoos  lost  animals  through 
this  cause,  but  now,  through  being  very  careful  not  to 
give  any  rough  spined  grasses,  we  never  have  the 
trouble."  Since  receiving  this  letter  we  have  re- 
moved straw  bedding,  and  feed  only  soft  alfalfa  which  is 
carefully  inspected  for  foreign  substances.  During  this 
time  we  have  had  two  cases  but  the  period  of  observation 
is  too  short  for  final  judgment  as  to  the  value  of  feeding 
grasses  without  sharp  beards  and  spines.  The  appear- 
ances of  cases  sporadically  without  reference  to  the  arri- 
val of  new  specimens,  the  low  communicability  of  the 


THE  STREPTOTHRICOSES  573 

disease,  its  beginning  in  the  jaw  in  most  cases  and  the 
prominence  of  pulmonary  and  gastric  lesions,  all  seem 
to  support  the  thought  that  the  virus  is  received  with  the 
fodder  or  drink.  The  anatomy  of  the  kangaroo's  buccal 
cavity  favors  the  collection  of  material  between  the  gums 
and  cheeks  and  between  the  root  of  the  tongue  and  the 
molars,  while  the  '*  hare  lip  "  also  affords  a  crevice  in 
which  food  particles  or  foreign  bodies  may  accumulate. 
These  three  places  seem  to  be  the  starting  points  of 
most  of  the  cases. 

The  Course  of  the  Attack. 

Despite  careful  watching  of  the  exhibition  specimens 
it  is  often  difficult  to  detect  the  beginning  stages  of  the 
disease.  Since  our  last  outbreak  it  has  been  the  practice 
to  examine  all  kangaroos  thrice  yearly  by  catching  them, 
inspecting  the  buccal  membranes,  teeth,  tongue  and  nose 
and  by  palpating  the  jugular  and  sublingual  regions.  This 
procedure  succeeded  in  catching  one  very  early  case  from 
which  the  original  changes  can  be  described. 

The  animal  appeared  in  generally  good  condition  but 
close  inspection  revealed  a  ''  running  nose,"  a  purplish 
mottling  along  the  gingival  margin  of  one  lower  jaw  be- 
low which  was  a  doughy  swelling ;  no  internal  ulceration 
had  appeared  nor  was  there  a  \'isible  change  in  the  exter- 
nal contour  of  the  jaw.  Within  a  few  days  a  small  fusi- 
form lump  appeared  along  the  body  of  the  lower  maxilla 
which  spread  gradually  backward,  the  nearby  soft  parts 
becoming  involved  very  shortly.  This  particular  animal 
died  without  ulceration  but  with  evidences  of  septicemia. 
Usually  at  the  time  that  the  lump  is  noticeable  the  animal 
loses  appetite,  becomes  inactive  and  seems  depressed;  no 
especial  change  in  the  coat  need  be  perceptible  although 
it  may  be  lusterless  or  at  times  ruffled.  In  the  cases  with 
great  involvement  of  the  cervical  tissues,  dyspnoea  is  an 
early  sign  but  I  lay  this  more  to  pulmonary  disease  than 
to  mechanical  obstruction  of  the  upper  air  passages.  The 


574  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

loss  of  appetite  is  in  large  part  due  no  doubt  to  the  dis- 
comfort of  chewing  and  swallowing  in  the  presence  of  an 
inflammatory  mass  in  the  neck.  The  eyes  usually  remain 
normal  until  quite  late.  No  change  in  the  character  of 
the  droppings  is  recorded. 

A  slightly  different  course  is  followed  by  the  cases 
that  have  the  primary  lesion  in  the  ''  hare  lip  "  and  nose ; 
from  these  the  masses  along  the  jaw  may  be  entirely 
missing.  After  an  initial  stage  of  * '  running  nose  ' '  ^vith 
or  without  swelling  of  the  upper  lips  and  alae  of  the 
nose,  the  animal  rapidly  goes  down  hill,  with  dyspnoea, 
loss  of  flesh,  perhaps  loose  stools,  lusterless  eyes  and  a 
*'  dead  "  coat,  a  series  of  signs  indicative  of  a  septicemic 
state  not  pronounced  in  the  first  variety ;  any  form  of  this 
infection  may  however  present  course  and  pathology 
of  a  septicemic  character.  If  the  beast  live  long  enough 
ulceration  may  appear  on  the  upper  lips  or  a  large  area  of 
necrosis  between  them  may  be  discovered. 

There  have  been  in  our  series  two  cases,  believed  to  be 
due  to  the  same  virus,  which  gave  a  picture  of  septice- 
mia with  pulmonary  localization;  they  will  be  discussed 
in  detail  later  but  are  of  interest  here  because  they  were 
not  known  to  be  sick  until  the  day  before  their  death.  A 
similar  failure  to  evince  signs  of  sickness  is  found  in 
the  gastrointestinal  cases,  those  with  ulcerations  in  the 
stomach  and  perhaps  an  accompanying  catarrhal  intes- 
tinal inflammation.  At  most  the  report  will  be  that  the 
specimen  was  ' '  off  its  feed. ' ' 

The  signs  of  Kangaroo  disease  mth  exception  of  those 
applying  directly  to  the  nose  and  jaw  are  therefore  very 
vague  and  one  is  limited  to  observation  of  the  contour 
of  the  head  and  of  the  discharge  from  the  nostrils.  Be- 
cause of  the  indefinite  nature  of  the  earliest  changes, 
the  duration  of  the  disease  cannot  be  stated  with  accuracy 
but  from  the  time  that  the  swellings  are  perceptible  it  is 
not  very  protracted  if  no  treatment  be  given.  Some  cases 
die  in  four  or  five  days  wliile  others  may  last  up  to  three 


Fu..  62. — KANGAROO  STREPTOTHRICOSIS.  ULCERATIVE  AND  NECROTIZING  PROCESS  IN 
"HARE-LIP"  AND  IN  MUCOSA  AND  BONE  OF  ANTERIOR  PORTION  OF  HARD  PALATE;  TEETH 
HAVE  FALLEN   OUT  ON  LEFT  SIDE. 


THE  STREPTOTHRICOSES  575 

weeks  and  we  believe  that  two  of  our  cases  may  have  been 
existent  longer  than  that.  It  is  impossible  to  estimate  the 
duration  of  the  septicemic  and  gastric  forms  although 
the  latter,  judging  by  the  appearance  of  the  ulcers,  are 
believed  to  be  chronic.  We  believe  that  frequent  inspec- 
tion and  the  precautions  as  to  the  character  of  fodder  are 
the  only  special  hygienic  measures  indicated. 

The  iNcroENCE  of  the  Disease. 

Not  the  least  puzzling  character  of  the  disease  is  the 
variability  of  its  appearance.  There  have  been  groups 
of  cases  in  our  records ;  for  example  the  following  periods 
showed  several  while  the  intervening  years  lacked  them 
entirely— 1905,  1907-8,  1911-2,  and  1920-1.  The  second 
and  fourth  outbreaks  were  definitely  related  to  a  new 
arrival  but  the  records  do  not  show  that  such  was  the 
case  for  the  other  two.  Mr.  Joseph  who  supplied  us  with 
many  specimens,  tells  us  that  he  has  had  an  experience 
of  fifty-four  cases  in  200  kangaroos  and  then  failed  to 
encounter  the  disease  for  years.  Perhaps  this  irregu- 
larity of  appearance  has  something  to  do  with  the  charac- 
ter of  food  supplied  to  the  animals. 

Among  seventy  deaths  of  Macropodidse  we  have  had 
thirty-three  cases  of  the  varieties  which  I  have  included 
in  this  infection,  made  up  of  the  following  forms :  cases 
limited  to  the  jaws,  pharynx  and  neck,  six;  cases  of  this 
sort  with  extension  to  lungs  and  stomach,  ten;  cases  of 
this  sort  with  general  spread  suggesting  septicemia,  five; 
gastrointestinal  and  hepatic,  eight;  nasal  and  sinus  in- 
fection \\dthout  necrosis  in  the  jaw  and  with  general 
spread  including  the  lungs,  four.  The  total  incidence  in 
Kangaroos  is  therefore  47  per  cent.,  the  necrotizing  forms 
being  30  per  cent.,  the  gastrointestinal  11.4  per  cent. 

The  Pathology. 

The  essential  features  of  the  necrotizing  variety  of 
this  disease  are  similar  to  those  of  actinomycosis — an  in- 
flammation giving  rise  to  much  fibrous  tissue  overgrowth 


576   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

enclosing  pockets  of  softening,  the  whole  process  causing 
a  deforming  tumefaction.  Wliile  primarily  developing 
in  the  soft  parts,  this  streptothricosis  behaves  like  the  ray- 
fungus  in  that  it  spreads  not  only  along  clefts  of  tissue 
but  directly  through  muscles  and  organs  and  even  bones. 
Rarefjdng  osteitis  with  irregular  attempt  at  repair  in  the 
f oral  of  productive  periostitis  may  be  found  in  both  infec- 
tions. There  is  however  a  greater  tendency  to  ulceration 
and  general  disease  in  the  marsupial  form,  variations 
which  seem  referable  to  secondary  invaders.  When  how- 
ever the  massive  tumorous,  necrotic  and  ulcerative  char- 
acters of  this  streptothrix  disease  are  insignificant  or 
absent,  the  pathology  is  modified  to  the  extent  of  obvious 
bacterial  mixed  infection,  there  then  being  catarrhal  and 
fibrinous  inflammations  mth  degenerations  of  the  viscera. 

Bacteriology. 

Since  the  pathology  varies  with  the  bacteriology  as 
seen  at  this  laboratory,  it  is  well  to  pause  at  this  time 
in  a  discussion  of  the  former  subject  to  introduce  a  brief 
statement  of  our  findings  in  the  latter,  leaving  however  a 
full  description  thereof  for  later  paragraphs.  It  is  rela- 
tively easy  to  find  in  smears  from  necrotic  masses  threads 
of  streptothrix,  straight  or  curved  with  heavy  blunt,  but 
not  bulbous,  ends  and  never  branched.  Similar  forms  may 
be  found  in  the  necrotic  tissue,  both  free  in  the  softened 
area  and  near  the  margin  of  the  healthy  tissue,  as  irregu- 
lar colonies  growing  in  a  tangled  mass  from  the  edge  of 
which  radiating  threads  may  be  seen.  It  has  not  been 
possible  to  find  a  ''  ray  "  growth  with  anything  like  the 
regularity  so  characteristic  of  the  actinomyces  nor  do 
the  ends  present  the  bulb  distinctive  of  that  organism. 
Branching  has  been  found  once  only,  it  being  very  un- 
common in  tissues  although  beautifully  developed  in  cul- 
tures. While  not  especially  sought,  cocci  and  bacilli  have 
not  been  seen,  by  Gram  stains,  within  or  immediately 
around  the  streptothrix  colonies.    Surrounding  the  myce- 


THE  STREPTOTHRICOSES  577 

lial  groups  is  a  necrotic  zone  about  which  is  a  loose 
connective  tissue  full  of  mononuclears  and  a  few  polynu- 
clears.  The  centre  of  the  colony  is  made  up  of  tangled 
mycelial  threads  and  necrotic  debris.  From  uncontami- 
nated  necrotizing  masses  we  have  obtained  cultures  three 
times  out  of  very  many  attempts. 

When  ulceration  or  suppurative  softening  has  taken 
place  mixed  infection  with  lower  bacterial  forms  natu- 
rally occurs  and  the  whole  picture  changes.  Pulmonary 
complications,  with  or  without  evident  ulceration  in  the 
pharynx,  also  admit  other  bacteria.  Streptothrical  forms 
are  often  easy  to  detect  in  stained  smears  and  in  cultures 
but  the  very  extensive  bacterial  flora  soon  overgrows 
them  and  attempts  at  isolation  are  fruitless.  Under  the 
best  of  conditions  their  colonial  development  is  slow 
and  tiny  until  they  are  well  accustomed  to  saprophytic 
life.  The  complicating  bacteria  that  have  been  identified 
are  Streptococcus  pyogenes,  pneumococcus,  pyocyaneus 
and  colon  bacilli  to  which  may  be  added  moulds  of  the 
Aspergillus  group  but  these  all  have  been  variable  in 
numbers  and  appearance;  the  most  frequent  and  there- 
fore probably  most  important  secondary  invader  is  an 
organism  we  have  not  been  able  to  identify. 

This  germ,  a  tiny.  Gram-negative,  non-motile  rod  with 
a  tendency  to  bipolar  staining,  will  appear  in  smears 
from  an  ulcerated  necrotic  mass,  from  the  nasopharyn- 
geal exudate  and  from  pulmonary  lesions  and  may  develop 
upon  agar  or  blood  media  for  the  first  generation  but 
refuses  to  grow  after  that  despite  our  best  efforts.  At 
present  we  hope  to  have  it  by  gTOwing  material  a  long 
time  in  blood  broth.  Microscopical  examination  has  not 
revealed  it  in  the  tumor-forming  variety  but  on  one 
occasion  it  was  found  in  the  lung ;  its  Gram-negative  char- 
acters make  its  detection  in  tissue  very  difficult.  For 
obvious  reasons  the  importance  of  this  germ  cannot  be 
estimated  but  it  seems  from  the  frequency  with  which 
it  is  encountered  that  in  some  manner  the  streptothrix 


578  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

may  be  aided  by  this  unidentified  bacillus  especially 
in  the  ulcerative  and  septicemic  varieties  of  Kangaroo 
disease. 

In  so  far  as  the  diagnosis  of  this  infection  in  the 
uncomplicated  form,  like  lumpy  jaw,  is  concerned  the  find- 
ing of  streptothrix  by  stain  seems  adequate  and  its  pres- 
ence in  the  gastric  ulcers  and  hepatic  necroses  identifies 
this  variety.  The  most  difiicult  question  to  decide  is  the 
identity  of  the  cases  without  one  or  the  other  of  these  dis- 
tinctive features  but  with  mucocatarrhal  or  purulent 
nasosinusitis  followed  by  pneumonia  or  septicemia,  and 
of  cases  of  primary  pulmonary  involvement.  These  in- 
stances have  been  diagnosed  as  belonging  to  the  same 
category  because  of  the  presence  of  streptothrix  in  the 
exudate  at  the  site  of  the  important  lesions  and  because 
the  type  of  lesion  is  similar  to  that  which  complicates 
accepted  characteristic  cases.  Inability  to  reproduce  ex- 
perimentally any  of  these  infections  limits  our  criteria 
for  judgment  in  the  matter.  I  am  inclined  to  view  these 
septicemic  cases  therefore  as  initiated  by  the  strepto- 
thrix, growing  in  the  nose  and  sinuses  or  inhaled  into  the 
lungs,  aided  by  lower  bacteria,  an  unidentified  Gram- 
negative  bacillus  being  the  most  important. 

Having  reviewed  briefly  the  bacteriology  of  Kangaroo 
disease,  its  strict  pathology  may  be  discussed  more  defi- 
nitely in  terms  of  the  type  of  infection.  Reference  has 
already  been  made  to  the  method  of  pathogenesis 
employed  by  the  streptothrix  and  its  congeners.  Whether 
or  not  a  toxin  is  elaborated  by  these  organisms  is  an 
unsettled  question,  especially  for  the  marsupial  variety 
because  as  yet  it  cannot  be  made  to  produce  lesions  in 
other  animals.  It  is  highly  probable  that  all  these  organ- 
isms find  colonization  easy  in  the  animal's  body  once  they 
get  well  settled,  and  that  they  act  mechanically,  producing 
necroses  by  their  gro^^i•h  and  by  attracting  leucocytes  in 
such  large  numbers  that  digestion  of  devitalized  tissue 
occurs,  to  an  extent  that  resembles  pus.    The  inflamma- 


Fig.    64. — KANGAROO     STRKPTOTHRICOSIS.       STOMACH.     SHOWING   TWO    ULCERATIONS    AND    DEEP 
INKU^TRATIONS   OF  THE  WALLS. 


THE  STREPTOTHRICOSES  579 

tory  tissue  is  not  distinctive,  except  in  so  far  that  fibrosis 
enclosing  pus  pockets  is  peculiar  to  it.  In  softer  tissue, 
like  the  liver,  fibrosis  is  not  so  prominent,  whereas  diffuse 
and  irregular  spread  is  more  pronounced.  At  the  margin 
with  the  healthy  tissue,  reactive,  that  is  resistant,  inflam- 
mation is  no  more  in  evidence  than  within  the  tumor 
growth  itself  and  as  a  matter  of  fact  the  tissues  do  not 
seem  to  put  up  a  good  fight  against  the  spread  of 
the  inflammation. 

Histologically,  aside  from  the  finding  of  the  strepto- 
thrix  colonies,  there  is  nothing  distinctive,  the  peculiar 
expressions  of  the  disease  being  most  manifest  in  their 
gross  characters.  For  the  purpose  of  describing  the 
pathological  features,  the  cases  have  been  divided  into 
the  necrotizing  form  around  the  jaw,  a  similar  process 
in  the  stomach  and  liver,  necrotic  cervical  cases  followed 
by  lung  involvement,  the  nasal  variety  upon  which  pneu- 
monia succeeds  and  a  septicemic  form  arising  from  any 
locality.  Illustrative  cases  will  be  cited  for  each  of  these 
forms,  a  method  of  presenting  the  pathology  thought  to 
be  superior  to  a  general  discussion. 

The  first  illustrative  case  is  one  localized  in  the 
tongue  and  pharyngeal  wall;  it  is  quoted  because  of  its 
strict  localization. 

Great  Gray  Kangaroo  {Macropus  giganteus) .  Sick  four  days,  tongue 
swollen  so  he  could  not  eat. 

Diagnosis. — Necrotizing  process  of  floor  of  mouth  and  pharyngeal 
wall,  dilatation  of  heart,  passive  congestion  of  liver,  acute  diffuse 
nephritis,  inflammatory  edema  of  lungs.  General  condition  good.  Jaws 
and  teeth  negative.  Floor  of  mouth  firm  in  places,  boggy  in  others, 
but  generally  infiltrated.  Anterior  two-thirds  of  tongue  purple  and 
green  as  if  gangrenous.  Root  of  tongue  and  adjacent  floor  of  mouth 
yellowish,  wet  as  if  from  recent  coagulation  necrosis.  In  the  muscle 
of  the  tongue  a  line  of  demarkation  is  shown  at  end  of  hemon'hagic 
zone  behind  which  muscle  is  fairly  good.  Sides  of  pharynx,  palate, 
tonsillar  region  show  superficial  pseudomembranous  inflammation  and 
yellowish  gray,  wet  infiltration  of  muscles.  Epiglottis  purple  and 
swollen  to  twice  normal  size.  Laryngeal  mucosa  deeply  injected,  swol- 
len and  covered  with  tenacious  gray  mucus.  Trachea  and  bronchi  deeply 
injected  and  slimy.    The  lungs  are  uniformly  deeply  injected  and  along 


580   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

course  of  bronchi  in  lower  lobe,  lung  tissue  is  distinctly  more  boggy  than 
elsewhere.  On  section  this  area  is  slightly  paler  and  more  granular  than 
the  rest  of  the  lung.  Lung  is  everj-^where  slightly  edematous.  The 
bronchial  lymphatics  are  swollen,  pale  pink  and  edematous.  The  heart 
is  dilated  acutely  judging  from  the  left  ventricle  wall  Avhich  is  nowhere 
over  1  cm.  The  liver  is  slightly  enlarged,  surface  smooth,  edges  sharj), 
color  deep  purple,  section  surface  very  bloody.  The  spleen  is  soft, 
capsule  smooth,  pulp  homogeneous  purple,  follicles  not  visible,  trabecu- 
lae  normal.  The  kidney  is  slightly  large,  capsule  smooth,  strips  easily 
leaving  purple  surface.  The  cut  surface  swells  out,  has  irregular  striae, 
congested  lines  betAveen,  glomeruli  visible  and  large.  Smear  from  centre 
of  tongue  muscle  shows  staphylococci  in  some  places  in  colonies,  and 
long,  slender  rods. 

The  foUoAviiig  case  is  one  of  gastric,  intestinal  and 
hepatic  involvement,  apparently  primary,  the  last  pos- 
sibly arising  by  a  hematogenic  or  lymphogenic  route. 
Judging  by  the  slides  of  the  gastric  wall  the  process 
started  deeply  and  broke  through  the  mucosa.  This  can- 
not be  asserted  definitely  since  kangaroos  are  susceptible 
to  gastritis  so  that  the  streptothrix  may  have  been 
implanted  upon  a  preexisting  inflammation. 

Black  Wallaby  {Macropus  ualabatiis).  Congestion  and  edema  of 
lungs,  abscess  of  stomach  and  liver  (streptothrix),  ulcerative  enteritis, 
necroses  of  spleen  and  lymph  nodes,  congestion  of  kidney.  The  animal 
is  thin,  hair  loose.  The  mouth  and  nose  seem  to  be  absolutely  healthy. 
The  weight  of  the  lungs  is  increased  by  congestion,  they  are  solid, 
homogeneously  red,  with  no  air  in  any  lobe  except  at  edges.  A  piece 
cut  from  centre  of  lung  sinks  quickly  in  water.  The  trachea  contains 
frothy  blood.  The  heart  muscle  is  soft,  flabby  and  lustreless,  chambers 
dilated,  valves  normal.  The  liver  is  of  normal  size,  firm,  smooth  sur- 
face, sharp  edges,  red  brown  color.  The  small  sublobe  of  the  liver 
which  lies  between  the  gall-bladder  and  the  pyloric  end  of  the  stomach 
shows  a  large  abscess  4x3  em.,  apparently  starting  in  the  substance 
of  the  liver  via  the  bile  ducts.  This  is  certainly  not  extension  from 
the  stomach  abscess  as  the  liver  lying  against  the  stomach  is  nearly 
normal.  The  abscess  is  sharply  circumscribed  with  a  zone  of  conges- 
tion about  it.  Aside  from  congestion  the  rest  of  the  liver  is  normal. 
The  common  bile  duct  is  large  and  freely  patulous.  The  capsule  of  the 
spleen  is  thick,  consistency  firm,  pulp  deep  red,  irregularly  mottled  by 
pale  areas  of  necrosis.  The  kidney  capsule  is  smooth,  strips  easily 
leaving  a  smooth,  brown  surface.  The  organ  is  firm.  The  section  sur- 
face is  glistening,  the  cortex  wide  and  congested,  the  medulla  normal. 
The  adrenal  medulla  is  deep  purple  A^dth  congested  line  between  it  and 
the  pale  cortex.     Most  of  the  gastric  mucosa  seems  good.     At  about 


THE  STREPTOTHRICOSES  581 

the  middle  of  the  lesser  curvature  is  an  ulcer  about  4  em.  across.  The 
shelving  edges  are  covered  mth  apparently  normal  mucosa.  The  centre 
contains  bloody  pus  and  nodular  masses  of  the  submucosa  extending 
in  finger-like  projections  through  the  pus.  At  one  point  on  the  greater 
curvature  there  is  a  small  pocket  of  pus  on  the  serous  side  which  has 
not  ulcerated  through  to  the  mucosa  nor  broken  into  the  peritoneum. 
The  large  intestine  is  deep  red  and  the  follicles  appear  from  the  serosa 
as  darker  areas.  On  the  mucus  side  the  follicles  have  ulcerated,  having 
a  necrotic  centre  and  shelving  edges.  The  rest  of  the  mucosa  in  the 
neighborhood  is  swollen  and  deep  red.  The  colon  mucosa  is  dry  and 
the  contents  are  hard,  dry  "  baked  "  feces.  The  main  pancreatic  duct 
and  the  common  bile  duct  form  a  thick,  firm,  cord-like  mass  running 
through  the  pancreas  and  enlarging  the  papilla  of  Vater  into  the  duo- 
denum. All  abdominal  lymph  nodes  are  large,  firm  and  on  section 
mottled  with  red  areas.  Culture  from  the  liver  abscess  failed  to  grow. 
Histological  section  of  lung  shows  moderate  congestion,  collapse  of 
alveoli  or  their  filling  by  edema,  epithelial  and  small  round  cells.  There 
seems  to  be  no  fibrin.'  This  could  be  an  early  stage  of  pneumonia. 
Bronchi  are  for  the  most  part  negative,  little  peribronchial  round  cell 
infiltration.  No  streptothrix  in  two  areas  of  round  cell  infiltration  or 
in  bronchi.  Liver  section  shows  a  part  of  the  liver  destroyed  by  hemor- 
rhage, degeneration  and  necrosis.  The  abscess  consists  of  necrotic  matter 
suiTOunded  by  a  zone  of  about  equal  numbers  of  mono-  and  polynu- 
clears  and  around  this  a  loose  fibrocellular  zone.  Streptothrix  abundant 
in  the  abscess.  Lymph  nodes  show  chronic  inflammation  and  coagula- 
tion necrosis  without  abscess  formation.  No  streptothrix  in  areas  of 
necrosis.  Kidney  is  very  much  congested  with  little  or  no  damage  to 
secreting  parts.  Spleen  shows  enormous  congestion,  moderate  amount 
of  pigmentation,  connective  tissue  both  trabecule  and  through  pulp 
increased,  no  areas  of  necrosis.  In  the  stomach  the  mucous  membrane 
shows  slight  cellular  activity  and  some  degeneration — this  amounts  to 
a  true  catarrhal  gastritis  especially  in  view  of  the  submucous  cellular 
infiltration  and  the  granulation  tissue  which  has  separated  the  muscu- 
laris  and  involved  most  of  the  connective  tissue.  The  edge  of  the 
necrotic  part  begins  abruptly,  the  mass  of  necrosis  lying  on  an  active 
fibrocellular  submucous  and  muscular  layer.  Streptothrix  can  be  seen 
at  edge  and  in  necrosis. 

Pneumonia  originating  either  by  inhalation  or  via  the 
blood  stream,  is  illustrated  in  two  stages  by  the  succeed- 
ing cases.  The  first  history  illustrates  the  pulmonary 
involvement  as  secondary  to  necrotic  streptothricosis 
around  the  jaw  and  tongue  while  the  second  animal's 
disease  began  in  the  nose  and  related  sinuses.  These 
two  protocols  provide  material  for  a  discussion  of  two 
phases  of  the  subject. 


582   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

The  character  of  the  early  bronchopneumonia  in  the 
first  is  peribroncliial,  and  there  is  distinct  indication  of  a 
generalized  process  suggesting  a  hematogenic  origin, 
whereas  there  is  but  one  area  of  bronchopneumonia  in  the 
second — a  necrotizing  lesion  beginning  in  the  bronchus. 
Streptothrices  are  rare  in  the  first  case  but  reasonably- 
easy  to  find  in  the  second.  This  latter  is  one  of  the  cases 
which  seem  to  support  the  idea  that  nasosinusitis  may 
have  a  streptothrix  as  its  basis  in  the  absence  of  the  usual 
picture  of  necrotizing  ''lumpy  jaw."  These  cases  also 
indicate  that  pneumonia  may  originate  either  by  inhala- 
tion or  by  the  blood  stream,  and  that  perhaps  the  hepatic 
lesion  may  have  the  latter  origin.  There  have  been  two 
instances  of  necrotizing  periarthritis,  in  one  of  which  the 
threads  could  be  found.  This  also  suggests  that  spread 
through  the  blood  stream  can  occur,  possibly  in  this 
respect  to  places  where  previous  injury  prepares  for  the 
reception  of  the  organisms. 

Thigh-striped  Wallaby  {Macropus  thetidis).  Streptothricosis  of  soft 
tissues  of  jaAV.  Early  bronchopneumonia.  Acute  fermentative  gastri- 
tis. Acute  general  infiltrative  enteritis.  Cloudy  swelling  of  myocar- 
dium. The  general  condition  of  coat  and  of  nutrition  is  good.  The  jaws 
are  wide  and  the  maxilloeervical  region  full,  both  due  to  an  indurative 
inflammation  of  the  gums,  tongue,  floor  of  mouth  and  upper  cervical 
tissues.  At  either  side  of  the  tongue  and  running  around  body  of 
maxilla  both  sides,  the  inflammatory  tissue  becomes  softer  and  there 
is  an  area  about  one  inch  long  Avhere  it  is  soft,  gray  and  contains  yellow 
gray  bodies  in  a  grumous  matrix.  The  teeth  seem  sound  as  do  the  ex- 
ternal buccal  tissues.  The  nasopharynx  is  free  from  induration.  The 
bone  on  the  left  side  shows  a  periosteitis  with  involvement  of  the  super- 
ficial layers  of  bone,  Avhile  on  the  right  side  the  periosteum  is  swollen 
and  opaque  but  the  bone  is  free.  The  thyroid  is  imbedded  in  the  edema- 
tous infiltration  of  the  lower  cervical  tissues.  The  pleura;  are  free  of 
fluid  and  adhesions.  Lungs  are  collapsed,  uniformly  pink  somewhat 
emphysematous  at  places  but  give  the  impression  of  being  lumpy.  On 
palpation  numerous  nodular  areas  are  detected.  These  prove  to  be 
peribronchial  areas  of  gray-red  solidity  which  swell  out  on  section. 
The  bronchus  contains  a  gray  and  bloody  thick  mucoid  matter.  There 
is  distention  of  the  mesenteric  vessels  especially  near  the  enteric  inser- 
tion. The  liver  surface  is  smooth,  edges  very  sharp,  consistency  firm, 
tough,  resilient,  color  deep  red,  the  section  surface  is  glistening,  moist, 


AND 

NOCA 


wm^  .^;--*,  ;■*#.-'' 

Fir..  66— KANGAROO  STRKPTOTHRK  OM  v  >K  I  loN  OF  V'\^',  ,f  I',",\"  "^.V,  ,\  nVv\  «  v'^rissflF^ 
NECROSKS.  ONE  WITHIN  A  BRONCHIA,  (.\1  IN  SKMICON^OLIDA  1  I  I  >  IMI.M..\AR^  I  ISSUE. 
iRDIAI,     STRANDS    COUI.D    BE   FOUND    IN    BOTH    AREAS. 


THE  STREPTOTHRICOSES  583 

opaque,  architecture  probably  normal.  The  gall-bladder  is  distended 
with  viscid  green  bile;  the  common  duct  is  patulous.  The  spleen  has 
a  rough,  thin  capsule,  consistency  tough  and  resilient,  the  section  sur- 
face is  mottled  red  with  purple  points ;  on  section  tAvo  small,  pale  objects 
seem  to  be  squeezed  out.  The  kidney  capsule  is  smooth,  strips  easily 
leaving  a  smooth,  deep  red  surface,  the  consistency  is  soft, 
the  cortex  is  deep  red,  then  a  purple  line  between  it  and  the  red 
medulla,  striae  invisible.  The  stomach  contains  frothy  grayish  mush. 
The  mucosa  is  finely  mammillated,  deep  pink  until  the  last  third  when 
it  becomes  deep  red,  deeply  injected  and  somewhat  thickened.  The 
pylorus  is  closed.  Externally  the  gut  is  congested,  in  places  translu- 
cent but  for  most  part  seems  thickened  by  reddish  swelling  of  both 
external  layers  and  mucosa.  The  mucosa  is  granular  or  pebbly  with 
here  and  there  a  small  bloody  suffusion.  The  histological  section  of 
lung  shows  alveoli  open,  septa  relatively  thin  but  somewhat  congested, 
bronchi  mostly  open  and  connective  tissue  not  increased.  Some  few 
bronchi,  especially  the  larger,  show  a  slight  catarrhal  bronchitis  but 
mostly  an  infiltrative  peribronchitis.  The  nearby  veins  and  arteries 
show  the  most  striking  change,  there  being  in  nearly  all  of  them  a 
distinct  thrombosis  without  circumferential  pneumonia.  In  one  place  a 
distinct  peribronchial  pneumonitis  was  found.  The  kidney  shows  very 
marked  congestion  of  all  parts,  causing  compression,  cloudiness  and 
granularity  of  the  epithelium.  Glomeruli  and  connective  tissue  about 
normal.  The  intestinal  serosa  is  negative  save  for  congestion.  Sub- 
mucosa  is  densely  infiltrated  with  mononuclears,  some  in  definite  groups. 
Section  does  not  show  areas  mentioned  in  notes  but  these  could  be  accu- 
mulations of  cells  Avith  congestion.  No  streptothrix  forms.  Section 
from  the  infectious  focus  of  face  consists  of  active  granulation  tissue, 
densely  injected  and  filled  with  mononuclears  of  two  types,  one  the 
lymphoid  cell,  the  other  of  the  young  connective  tissue  type.  Areas  of 
grouping  like  abscesses  are  seen  and  some  necroses.  Streptothrix  in 
small  numbers  in  the  cellular  collections. 

Nail  tailed  Wallaby  {Macropus  unguifer).  Kangaroo  disease  of 
nasal  region.  Necrotizing  bronchopneumonia  (Aspergillus  fumigatus 
and  Micrococcus  albus).  Acute  diffuse  splenitis.  Congestion  of  liver 
and  kidney.  The  general  condition  of  coat  and  nutrition  is  good.  The 
face  is  wide  just  below  the  eyes.  About  the  "  hare  lip  "  and  the  nose 
the  soft  tissues  are  soft,  gray,  necrotic.  All  the  internal  nasal  tissues 
seem  swollen,  gray-red.  There  is  subcutaneous  edema,  bloody  in  places, 
around  the  right  face,  eye  and  jugular  angle.  Tissues  of  nasopharynx 
swollen,  deeply  injected  and  covered  by  a  thick  mucus.  Pharyngeal  and 
buccal  cavities  negative.  Tonsillar  areas  pink  and  flat.  Larynx  and 
trachea  slightly  swollen  but  pale  on  mucosa.  Salivary  glands  and  cervi- 
cal glands  normal  in  size  and  pale  pink.  Pleurae  pale  and  empty.  Lungs 
swollen  out  uniformly,  quite  cottony  except  at  lower  right  base  where 
there  is  a  nodule  about  3x5  cm.  firm  and  doughy.  On  section  it  is 
found  to  be  a  peribronchial  consolidation  of  pale  reddish  gray  color 


584   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

and  indefinite  outline.  The  bronchus  itself  is  deeply  congested  and  eon- 
tains  a  grumous  mass.  The  peribronchial  lymph  nodes  are  small,  soft, 
pink,  homogeneous.  The  heart  is  negative.  The  liver  is  large,  surface 
smooth,  edges  sharp,  color  deep  purple,  consistency  soft.  Section  surface 
is  glistening,  smooth,  moist,  very  dark  purple  with  obscure  markings. 
The  gall-bladder  contains  fluid  brown  bile;  common  duct  is  patu- 
lous. The  spleen  is  soft,  tough,  capsule  pebbly,  section  surface  is 
mottled,  light  and  deep  pink,  follicles  and  trabeculae  not  distinguished. 
The  kidney  capsule  is  smooth,  strips  easily  leaving  a  smooth  purple 
surface,  section  surface  is  glistening,  deeply  congested,  striae  obscure 
but  seem  normal,  glomenali  not  visible,  organ  is  soft.  The  gums  and 
teeth  are  not  involved  in  the  mycosis.  The  stomach  contains  mushy 
digesting  food.  The  mucosa  is  mottled  pink,  soft,  digesting,  at  lower 
half  submucosa  is  deep  pink,  a  few  small  ecchymoses.  From  pylorus  to 
ileum,  serosa  is  deeply  injected,  edematous,  mucosa  swollen  and  edema- 
tous, deep  pink,  loosened  in  places,  but  translucent.  Below  this  the 
mucous  membrane  becomes  smooth,  flat,  pink-yellow.  Lower  ileum  and 
colon  contain  rather  firm  fecal  balls.  Follicles  nowhere  prominent. 
The  pancreas  is  small,  soft,  yellow-pink.  The  follicles  of  the  mesentery 
are  small,  pink-gray  and  homogeneous.  Smears  from  the  broncho- 
pneumonia show  a  thread-like  Gram-positive  form  and  a  few  Gram- 
negative  rods.  Cultures  from  lung  show  Aspergillus  fumigatus  and 
Micrococcus  alhns.  Nose  too  foul  for  culture.  Histological  section  of 
lung  shows  the  alveoli  mostly  open  but  the  septa  widened  by  congestion. 
Blood  vessels  are  open  and  contain  recent  clots;  one  vessel  near  lesion 
below  is  thrombotic.  The  two  large  bronchi  in  section  show  catarrhal 
bronchitis  and  infiltrative  peribronchitis  of  which  the  latter  is  more 
severe  and  advanced.  Beside  the  larger  is  a  necrotizing  pneumonitis 
from  which  nearly  all  the  architecture  has  disappeared.  The  exudate 
is  chiefly  mononuclear  around  the  edges;  centre  no  cellular  identity. 
Another  mononuclear  process  not  connected  with  bronchus  in  section 
is  found  with  an  early  necrosis.  Streptothrix  strands  may  be  found  in 
the  bronchial  exudate  and  near  the  margin  of  the  necrotic  patch.  They 
do  not  grow  in  colonies  however.  The  spleen  shows  general  congestion 
without  pigmentation.  Follicles  large,  solidly  lymphoid.  Connective 
tissue  about  normal.  The  kidneys  show  marked  congestion  everywhere. 
Capsule  and  intrarenal  fibrous  tissues  about  normal.  Very  severe  con- 
gestion which  seems  to  have  caused  compression  and  granularity  of 
the  epithelium. 

The  last  case,  judging  by  stained  smears,  is  one  of 
pure  nasosinusitis  from  streptococci  and  streptothrices. 
Cultures  were  not  tried  because  of  the  enormous  bac- 
terial flora. 

Robust  Kangaroo  {Maeropus  robustus) .  Acute  purulent  ethmoiditis. 
General  acute  purulent  anterior  cranial  sinusitis.     Acute  necrotizing 


^^T  Jx',^  67— KANGAROO  STREPTOTHRICOSIS.  LOW  POWER  PHOTOMICROGRAPH  OFANOCARDIA 
COLONY  WITH  NECROSIS  WITHIN  AND  AROUND  IT.  THIS  WAS  FOUND  IN  A  SECTION  FROM 
THE  LIP  OF  THE  SPECIMEN  SHOWN  IN  FiG.  62.  THE  BLACK  BORDER  CONSISTS  OF  PARALLEL 
THREADS  SO  CLOSELY  PLACED  THAT  THEIR  SEPARATION  UNDER  THE  CAMERA  IS  PRACTICALLY 
IMPOSSIBLE.      THIS   TYPE    OF  COLONY   RESEMBLES  THE   "RAY"   COLONY  OF  ACTINOMYCES 


THE  STREPTOTHRICOSES  585 

glossitis  and  pharyngitis.  Cloudy  swelling  of  kidney.  The  face  seems 
a  little  full  and  the  subcutaneous  tissues  slightly  edematous.  The  naso- 
pharynx contains  a  thick  tenacious  mucopus.  Ethmoid  and  frontal 
sinuses  and  turbinate  spaces  contain  a  thick  purulent  matter,  the  mucosa 
being  densely  injected,  swollen  and  velvety.  Pharyngeal  wall  and  right 
half  of  posterior  half  of  tongue  are  involved  in  a  dull  brown  and  necro- 
tizing process,  quite  sharply  outlined  by  zone  of  congestion.  This 
process  is  comparable  to  the  necrotizing  gingivitis  seen  in  front  of  jaw 
in  kangaroos.  Larynx,  trachea  and  lungs  seem  uninvolved  save  for 
slight  generalized  congestion.  Cervical  lymph  nodes  especially  those 
about  the  larynx  are  definitely  enlarged,  soft,  moist  and  brown.  Medias- 
tinal nodes  slightly  enlarged,  soft  and  pink.  The  heart  is  negative. 
Liver  normal.  Spleen  is  soft,  homogeneous  dull  red.  The  capsule  of 
the  kidney  is  smooth,  strips  easily  leaving  a  purple  surface.  The 
glistening  section  surface  swells  slightly,  vasa  recta  are  congested,  striae 
wide  and  pale,  glomeruli  not  visible ;  consistency  is  resilient.  The  mouth 
and  teeth  are  not  involved  in  the  process  mentioned  above.  There  is 
a  small  quantity  of  properly  digesting  food  in  the  stomach.  Stomach 
and  intestines  negative.  Brain  not  involved.  No  extension  from  anterior 
cranial  sinusitis.  Smears  from  the  mucopus  confirm  the  gross  appear- 
ance and  contain  short  chains  of  streptococci  and  large  diplococci. 
Smear  from  cut  surface  of  tongue  shows  innumerable  small  bacUli  and 
diplococci  but  especially  mycelia  with  rather  heavy  clubbed  ends  but 
without  true  branching.  One  group  was  found  arranged  like  ray  fun- 
gus. It  is  noteworthy  that  there  is  no  aspiration  pneumonia  and  very 
slight  evidences  of  septicemia. 

Biology  of  N.  macropodidarum. 

The  original  discovery  of  the  streptothrical  forms  was 
made  in  stained  smears  from  necrotizing  lesions.  They 
were  considered  as  secondary  invaders  until  repeated 
observations  of  a  similar  character  aroused  the  suspicion 
that  they  stood  in  some  important  relationship  to  the 
lesion.  Early  attempts  at  their  cultivation  were  made 
under  anaerobic  precautions,  a  method  now  known  to  be 
almost  certainly  doomed  to  failure  because  a  strain  long 
under  cultivation  requires  two  to  three  weeks  to  make  an 
appreciable  growth  in  the  absence  of  air.  Finally  in  1911 
a  successful  cultivation  occurred  by  searing  the  surface  of 
an  unopened  mass  in  a  freshly  dead  animal  and  planting 
bits  of  the  interior  upon  aerobic  blood  serum  plates. 
Colonies  grew  after  three  or  four  days  and  from  them  the 
first  strain  was  started.  It  grew  for  several  generations, 
38 


586  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

long  enough  for  the  preparation  of  a  vaccine,  which  will 
be  described  later,  when  by  mischance  it  was  lost.  In 
1920  another  successful  cultivation  occurred,  this  time  by 
incising  a  mass  in  the  soft  sublingual  tissue  and  plating 
in  the  same  manner;  upon  this  culture  the  biology  is 
described.  Smear  preparations  offer  no  more  than  has 
already  been  mentioned. 

Colonies  develop  upon  blood  serum  plates  as  opaque, 
pale  yellow,  circular,  discrete  masses  with  a  slightly 
depressed  uneven  centre,  but  without  umbilication. 
They  remain  smooth  and  slightly  glistening  for  several 
days,  then  become  slightly  wrinkled  and  twisted  with  a 
more  definitely  raised  edge  and  a  tendency  to  an  uneven 
sinking  in  of  the  centre.  Transfers  to  agar  slants  show 
wrinkled  continous  opaque,  dull  yellow,  sharply  outlined 
growths  which  soon  wrinkle,  fold,  and  twist  like  certain 
tubercle  bacillus  cultures.  Spreading  occurs,  but  is  slow 
after  forty-eight  hours.  As  medium  becomes  drier  it  is 
possible  to  see  a  thin,  colorless,  wrinkled  film  stretching 
out  from  the  main  growth.  If  the  medium  be  dry  or  old 
or  if  only  a  small  portion  of  seed  material  be  used  and 
this  scattered  over  the  surface  of  the  slant,  discrete 
colonies  arise.  These  are  circular,  seldom  exceeding 
3  mm.,  dirty  yellow-white,  distinctly  umbilicated  and 
without  clear  film  of  spreading  around  them. 

In  nearly  all  quite  old  cultures,  a  white  chalky  efiSorescence  appears 
over  the  surface. 

The  morphology  of  the  young  agar  culture  is  chiefly  mycelial  or 
filamentous,  whereas  from  a  culture  on  dried  media  and  those  show- 
ing efflorescence,  the  organisms  are  short,  heavy,  deeply  granular  and 
of  the  mycobacterial  type. 

Glycerine  agar. — Corresponds  to  agar. 

Blood  agar. — Similar  to  agar  but  much  less  luxuriant. 

Blood  serum. — Limited  dirty  yellow,  raised,  dull,  wrinkled  and 
granular,  tightly  adherent  to  the  medium. 

Potato. — Spreading,  dirty  yellow,  much  wrinkled,  friable,  tightly 
adherent. 

Gelatine. — Limited  growth  as  a  wrinkled,  tough  scum  only 
on  surface. 


Fig.  68.— kangaroo  STREPTOTHRICOSIS.  HIGHER  MAGNIFICATION  OF  EDGE  OF  STREPTO- 
THRIX  COLONY,  FiG.  67.  IT  SHOWS  THE  DEEPLY  STAINING  MYCELIA  SEPARATING  MUSCLE 
FIBRES  WHICH  ARE  DEGENERATING. 


THE  STREPTOTHRICOSES  587 

Litmus  milk. — No  change  for  six  days,  then  beginning  slight  alka- 
linity which  increases  very  little,  shows  digestion  of  the  caseinogen, 
slight,  thin  filmy  growth  on  surface. 

On  media  such  as  litmus  lactose  agar  and  old  Endo  it  grows  slowly 
on  surface  and  assumes  the  color  of  the  medium. 

Broth. — Only  surface  growth  appearing  during  early  generations, 
after  3-6  days  as  a  wrinkled,  pale  yellow  scum  very  much  like  the 
tubercle  bacillus  growth;  later  generations  grow  perceptibly  in  one  to 
three  days.  Medium  perfectly  clear.  If  a  large  mass  be  seeded  into 
neutral  broth  there  is  a  perceptible  increase  in  the  growth  after  ten 
days.  The  medium  thereafter  tends  to  a  faint  turbidity.  Titration 
of  broth  growth  after  twelve  days  shows  alkalinity  requiring  0.3  cc. 
decinormal  acid,  while  the  control  tube  incubated  same  length  of  time 
showed  an  acidity  requiring  0.57  cc.  of  decinormal  NaOH. 

On  the  following  sugars  there  is  a  slight  surface  growth  without 
change  in  the  color,  Andrade  indicator — dextrose,  lactose,  saccharose, 
maltose,  mannite,  dextrin,  galactose,  salicin. 

Cultures  observed  on  two  per  cent,  neutral  agar. 
A. — Stained  by  Loeffler's  stain. 

Twenty-four  hours. — Shows  threads  growing  out  from  a  central 
amorphous  mass,  but  the  whole  does  not  retain  the  regularity  or 
parallelism  of  actinomyces.  Threads  are  poorly  stained  and  rather 
disconnected  but  not  jointed.  Small  number  of  metachromatic  bodies 
apparently  in  older  individuals,  certainly  in  the  better  formed  ones. 
No  intercalary  spores,  unless  the  metachromatic  bodies  be  so  con- 
sidered. Individual  threads  measure  from  one-third  to  one  micron  in 
width.    Metachromatic  bodies  measure  on  the  average  one  micron. 

The  threads  in  the  forty-eight  hour  preparation  seem  distinctly 
wider,  up  to  one  micron  and  possibly  become  heavier  toward  the  end, 
but  do  not  have  a  distinct  bulbous  extremity. 

In  three  days  the  threads  are  much  longer,  show  distinct  branch- 
ing and  a  tendency  to  transverse  segmentation.  More  than  one  meta- 
chromatic body  may  be  present  in  one  segment. 

Four  days. — Still  coarser,  short  segments  have  appeared  separately. 
Metachromatic  body  is  coarser  and  blacker;  some  of  the  masses  have 
gone  to  pieces  and  show  only  a  diffusely  staining  smudge  of  metachro- 
matic bodies.  The  short  segments  show  a  tendency  to  grow  out 
into  threads. 

Fifth  day. — Condition  is  much  the  same  plus  many  young,  delicate, 
poorly  staining  threads. 

Sixth  day.^ — The  same  but  all  seem  to  be  somewhat  wider  and 
diffusely  staining. 

Seventh   day. — More   diffuse   staining   and   decidedly   fewer  meta- 
chromatic bodies. 
A. — Stained  by  Gram's  stain. 

Twenty-four  hours. — All  forms  are  light  purplish.  The  threads 
stain  much  more  clearly  than  by  Loefifler's  and  show  distinct  transverse 


588   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

segmentation  of  rather  uniformly  long  bacilliform  shape.  Metachro- 
matic bodies  not  so  distinct  but  seem  larger  Avhere  found.  Coarser 
threads  have  swellings  iji  some  of  the  areas  which  are  not  segmented  and 
this  type  seems  to  have  more  branching  and  metachromatic  bodies;  in 
other  words  it  would  seem  that  this  is  a  form  that  reproduces  by  budding 
or  intercalary  spore  formation. 

Forty-eight  hours. — Much  the  same,  more  long  threads  with  trans- 
verse division,  somewhat  more  delicate,  generally  fewer  coarse  threads 
with  swellings  and  spores.  Still  pale  purple  and  not  distinctly  Gram- 
positive. 

Three  days. — ^Condition  much  the  same. 

Four  daj-s. — Two  forms  present — definitely  Gram-negative  delicate 
slender  threads,  nearly  Gram-positive,  and  heavier,  curved  and  twisted 
long  bacillary  forms,  some  streptococcoid  threads  and  a  few  bulbous 
short  threads.    Very  few  metachromatic  bodies. 

Five  days. — Condition  much  the  same  except  that  the  delicate 
threads  are  inconspicuous  and  the  darker  purple  bacilli  have  increased. 
Metachromatic  bodies  increased  as  have  swellings  in  coarser  threads. 

Six  days. — Much  the  same  but  for  the  appearance  of  young,  deli- 
cate definitely  Gram-negative  threads.  There  are  fewer  metachromatic 
bodies  and  internal  spores. 

Seven  days. — The  same. 
B. — Grown  on  Loeffler's  blood  serum. — Loeffler's  stain. 

Twenty-four  hours. — Delicate,  poorly  stained  short  threads,  few 
tiny  metachromatic  bodies. 

Two  days. — Not  well  stained,  relatively  short  threads  show  numer- 
ous metachromatic  bodies  varying  from  exceedingly  tiny  dots  to  coarse 
granules  wider  than  the  thread.  These  may  be  numerous  in  the  same 
segment  and  form  a  row  from  six  to  ten.    Many  short  bacillary  forms. 

Three  days. — Poorly  stained,  metachromatic  bodies  apparently  more 
numerous  but  much  smaller. 

Four  days. — Almost  entirely  short,  heavy  bacillary  forms,  some 
of  which  are  very  like  diphtheria  bacillus  in  the  irregularity  of  width; 
many  metachromatic  bodies,  distinct  branching,  some  of  the  small 
heavy  ones  have  fusiform  swellings ;  practically  no  long,  heavy  threads. 

Five  days. — Essentially  the  same,  individual  elements  slightly  larger, 
fewer  but  coarser  metachromatic  bodies,  more  numerous  round  forms 
suggesting  large  pale  cocci. 

Six  days. — Much  the  same  but  elements  shorter,  smaller  and  some 
more  segmented. 

Seven   days. — ^More  long   forms  of   uniform   staining  but  still   a 
majority  of  coccoid  or  short  bacillary  forms  with  irregular  staining 
and  metachromatic  bodies;  no  long  threads. 
B. — Gram's  stain. 

Twenty-four  hours. — Pale  purple,  almost  Gram-negative,  long,  slen- 
der but  well  outlined  threads,  a  few  coccoid  forms,  practically  no 
granules. 


-KANGAROO  STREPTOTHRICOSIS.    PHOTOMICROGRAPH  SHOWING  THE  SEPARATE  THREADS 
OF  NOCARDIA  IN   A  SOFT  NECROTIC  LESION. 


THE  STREPTOTHRICOSES  589 

Two  days. — Very  pale,  almost  Gram-negative  threads,  very  many 
eoccoid  forms  and  short  rods,  considerable  segmentation  of  the 
longer  threads. 

Three  days. — Increase  in  short,  heavy  bacillaiy  forms  with  bulbous 
ends,  deeply  stained  ones  and  the  granules  being  lightly  Gram-positive ; 
long,  slender  threads  are  disappearing. 

Four  days. — Almost  exclusively  short,  heavy  forms  with  bulbous 
ends  with  eoccoid  forms,  heavier  forms  almost  definitely  Gram-positive, 
granules  Gram-positive. 

Five  days. — Much  the  same  but  more  segmentation  in  the  bacillary 
forms,  eoccoid  forms  become  more  numerous. 

Six  days. — Individuals  are  somewhat  longer  but  there  are  many 
rods  with  fusiform  swellings  containing  granules;  eoccoid  forms  pres- 
ent in  chains  sometimes. 

Seven  days. — More  long  rods  or  short  threads,  pure  eoccoid  and 
bacillary  forms. 

The  morphology  upon  bouillon  depends  somewhat  on  age  and  upon 
the  location.  Upon  the  surface  the  long  branching  mycelial  type  appears 
early  and  persists  until  the  whole  surface  is  covered  whereupon  the 
segments  divide  into  eoccoid  elements  with  metachromatic  bodies.  If 
heaping-up  develop  the  coarse  grains  on  the  mass  consist  of  granular 
or  eoccoid  rods.  When  growing  in  the  depth  the  eoccoid  form  is  the 
predominant  one,  only  a  few  delicate  mycelia,  usually  Gram-negative, 
being  found. 

The  Gram  character  of  the  organism  should  be  emphasized.  The 
young,  delicate  mycelia  are  negative  or  take  a  very  feeble  blue  stain. 
The  hea\^  bacillary  forms  are  Gram-positive.  Like  the  ray  fungus 
the  heavy  ends  are  sharply  Gram-positive,  but  unlike  it,  there  has 
never  been  seen  a  Gram-negative  bulbous  capsule  around  this  end. 

The  determination  of  this  organism  was  undertaken 
from  the  classifications  of  Petruschky  (Kolle-Wasser- 
mann),  of  Castellani  in  Castellani  and  Chalmers' 
Tropical  Medicine,  and  of  the  Society  of  American  Bac- 
teriologists. In  the  first  classification  it  corresponds  in 
some  ways  with  Streptothrix  Jwminis,  and  in  some  ways 
with  Streptothrix  caprce.  As  for  the  second  authority  it 
falls  into  the  Nocardiacese,  section  parasitica,  subsec- 
tion I,  in  that  a  distinct  earthy  odor  is  absent  and  that 
there  is  no  liquefaction  of  coagulated  protein.  It 
resembles  several  of  the  species  given  in  this  subsection, 
but  does  not  correspond  exactly  with  any  of  them.    Con- 


590   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

sultation  of  the  classification  of  the  American  Bac- 
teriologists would  place  it  among  Mycobacteriaceae.  The 
facts  that  it  is  strongly  aerobic,  produces  whitish  efflo- 
rescence which  may  possibly  be  aerial  hyphae  and  breaks 
up  into  short  segments,  place  it  in  the  genus  Nocardia. 
It  seems,  however,  to  belong  to  a  division  of  Nocardia 
which  is  close  to  the  Mycobacterium  since  the  short  ele- 
ments are  swollen,  cuneate  and  usually  heavy,  wliich  is 
imusual  in  the  more  typical  Nocardia.  It  is  not,  however, 
acid  fast  and  therefore  cannot  be  classified  among  the 
Mycobacteria.  This  culture  seems  to  be  a  variety  not 
heretofore  described,  and  since  its  association  with  the 
disease  is  so  definite,  whether  or  not  it  be  the  cause,  the 
name  Nocardia  MACROPODroAEUM  is  proposed,  because 
the  kangaroos  belong  to  the  order  Marsupialia, 
family  Macropodidae. 

The  discovery  of  these  organisms  within  tissues  is  by 
no  means  easy  even  though  the  larger  colonies  may  be 
located  by  staining.  If  Loeffler's  method  be  used  the 
central  mass  stains  quite  diffusely  and  the  spreading 
mycelia  around  the  edge  stain  faintly.  For  study  pur- 
poses this  stain  is  preferable  to  Gram-Weigert,  since 
despite  the  positivity  of  the  cultures,  the  blue  dye  can  be 
removed  very  easily  from  sections  and  only  with  great 
care  will  enough  remain  to  permit  tracing  of  the  separate 
threads ;  with  Gram  stain  no  detail  can  be  made  out  in  the 
centre  of  the  colony,  it  being  a  diffuse  blue.  Careful 
search  near  the  edge  of  these  necroses  will  usually  succeed 
in  the  discovery  of  a  few  mycelia  stretching  in  between 
the  mono-  and  polynuclears  of  the  low  grade  inflammation. 
This  is  best  seen  in  the  margin  of  gastric  ulcers,  but  may 
also  be  found  in  the  cervical  masses.  When  searching  in 
the  pulmonary  tissues  the  organisms  are  to  be  found  in 
the  bronchial  exudate  or  at  the  edge  of  pneumonias.  In 
one  nasal  mucosa  the  mycelia  were  dispersed,  not  gromng 
in  colonies  as  in  localized  inflammations. 


THE  STREPTOTHRICOSES  591 

Experiments  at  the  Repeoduction  of  the  Disease. 

When  the  first  culture  was  isolated  it  was  injected  into 
gTiinea-pigs ;  its  loss  stopped  further  work  because  it 
could  not  be  regained  from  the  animals.  The  present 
culture  had  been  injected  into  g-uinea-pigs,  rabbits,  opos- 
sums— all  with  negative  results;  such  an  experience  is 
not  unknown  for  actinomyces.  Intraperitoneal,  intra- 
venous methods  having  failed,  inoculation  was  made  into 
the  gums  of  rabbits  and  of  opossums  with  no  result,  even 
after  traumatizing  the  mucous  membrane.  The  injection 
of  about  5.  mg.  of  a  twenty-four-hour  agar  culture  was 
made  directly  into  the  masseter  muscle  of  an  opossum 
without  producing  even  a  lump  at  the  site.  Atomizing  a 
culture  into  the  nose  and  throat  of  an  opossum  seemed 
also  without  effect.  Injection  of  cultures  into  the  nose, 
gums  and  labial  tissues  of  a  wallaby  have  been  negative ; 
nor  has  any  perceptible  effect  followed  the  atomizing  of 
a  heavy  nocardial  suspension  in  broth  into  the  trachea  of 
this  animal. 

The  results  of  these  experiments  are  in  accord  with 
those  of  many  similar  attempts  to  reproduce  actinomy- 
cosis. Perhaps  in  Kangaroo  disease  the  small  Gram- 
negative  bacillus  is  a  necessary  factor. 

Specific  Prevention  and  Treatment. 

Encouragement  that  we  were  upon  the  right  track  was, 
however,  found  in  another  direction.  Improvement  in 
human  and  bovine  actinomycosis  having  followed  the  use 
of  vaccines,  it  occurred  to  me  to  try  this  method  as  treat- 
ment and  prophylaxis.  The  first  culture  to  be  isolated 
was  just  at  hand,  so  that  it  could  be  used  at  once.  Five 
injections  were  given  under  the  skin  of  the  thigh  to  a 
recently  developed  case  of  the  ulcerative  gingival  variety, 
a  noticeable  improvement  occurring  almost  at  once,  and 
at  death  there  was  an  apparent  cure  of  the  local  lesion. 
However,  the  accompanying  protocol  made  at  the  time 


592   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

tells  the  whole   story,  no   adequate   explanation  being 
at  hand. 

Red  Kangaroo  [Macropus  rufus).  Disease  of  the  mouth  first 
noticed  March  31,  1912,  died  September  13,  1912.  Necrotizing  osteitis, 
arthritis  and  periarthritis  of  left  ankle,  subacute  fibrinous  right  pleuri- 
tis,  hemorrhagic  bronchitis  with  atelectasis  in  right  middle  lobe,  abscess 
of  right  middle  lobe;  passive  congestion  of  lungs,  liver,  kidney,  chronic 
splenitis,  chronic  general  lymphadenitis.  The  animal  is  in  general 
good  condition  except  for  a  fusiform  swelling  about  the  left  heel  with 
evidence  of  fracture.  The  necrotic  process  in  the  hare  lip,  nose  and 
palate  has  entirely  disappeared.  One  front  incisor  has  gone  and  the 
other  is  loose.  There  is  a  scar  on  the  under  part  of  the  soft  palate 
in  a  small  healed  channel  between  palate  and  floor  of  nose.  There 
is  no  evidence  of  pyorrhoea.  Cervical  and  axillary  lymph  nodes  are 
much  enlarged,  pale  yellow,  firm  and  of  the  appearance  like  early 
stages  of  Hodgkin's  disease.  Faseias  of  cavities  congested.  The  lungs 
are  mottled  purple,  air  content  decreased,  section  surface  purple, 
exuding  frothy  blood.  The  whole  right  lung  is  covered  with  a  thick 
fibrinous  exudate,  most  intense  over  middle  lobe  at  site  of  atelectasis. 
There  are  light  scattered  adhesions.  The  anterior  margin  of  the  lung 
is  adherent  to  the  pericardium  which  is  covered  in  the  front  by  exudate. 
Upper  and  lower  lobes  show  hypostatic  congestion.  Middle  lobe  has 
separate  bronchus  filled  Avith  necrotizing  blood  clot  extending  into  a 
smaller  bronchus  with  complete  occlusion.  The  alveoli  supplied  by  the 
last  show  atelectasis  like  hemorrhagic  infarct.  There  is  a  small  sub- 
pleural  abscess  near  the  margin  of  this  atelectatic  area.  The  bronchial 
lymph  nodes  are  slightly  enlarged,  mottled  yellow  and  pink,  firm  with 
large,  diffuse  follicles.  The  pericardium  contains  2-3  cc.  clear  fluid. 
The  heart  muscle  is  pale,  purple  and  soft.  All  the  vessels  are  full  of 
currant  jelly  clot.  On  the  posterior  surface  of  the  aorta  internally  about 
an  inch  above  the  valves  there  is  a  patch  of  roughening  with  a  sugges- 
tion of  thickening  and  opacity.  It  is  comparable  to  the  early  stages 
of  syphilitic  aortitis.  The  liver  is  noiTual  in  size,  surface  smooth,  edges 
sharp,  consistency  firm  and  friable,  color  purple.  The  section  surface 
is  glistening,  smooth,  moist,  and  shows  passive  congestion.  The  gall- 
bladder contains  fluid  brown  bile  and  the  common  duct  is  patulous. 
The  spleen  is  slightly  enlarged,  firm  and  tough,  capsule  Avrinkled.  Sec- 
tion surface  is  mottled  red  and  purple  with  irregular  gray  trabeculoe 
and  faint  scattered  follicles  Avith  diffuse  margins.  The  kidney  capsule 
is  smooth,  strips  easily  leaving  a  smooth  brown  surface.  Organ  is 
firm.  The  section  surface  is  glistening  with  a  line  of  passive  congestion 
with  distended  vessels  between  the  cortex  and  medulla  which  are  of 
noi-mal  -n-idths.  Intestines  seem  normal  throughout.  The  pancreas  is 
firm,  pale  pink,  slightly  edematous.  The  mesenteric  lymph  glands  are 
moderately  enlarged,  yellow,  firm,  homogeneous  Avith  congested  centres. 
About  the  left  ankle  joint  there  is  a  necrotizing  infection  Avhieh  has 


THE  STREPTOTHRICOSES  593 

involved  the  bone  causing  a  pathological  fracture  of  the  lower  end  of 
the  tibia.  Smears  from  the  periarthritis,  pleuritis  and  blood  clot  in 
the  bronchus  show  streptothrix,  a  short  colon-like  rod  and  a  coccus  in 
fours — a  picture  precisely  like  that  obtained  from  the  jaw  bone  cases. 
In  addition  to  the  above  there  is  a  very  distinct  encapsulated  pneurao- 
coecus  form  in  smears  from  the  blood  clot  in  the  bronchus.  This  is 
the  animal  which  was  vaccinated  with  a  culture  made  from  the  depths 
of  a  necrotic  mass,  upon  which  treatment  she  rapidly  improved  and 
as  seen  from  the  above  notes  recovered  from  the  palate  condition.  Why 
she  should  have  a  second  infection  apparently  with  the  same  organism 
is  difficult  to  determine.  Possibly  the  second  batch  of  vaccine  was  not 
sterile,  it  not  having  been  controlled  because  the  fii'st  batch  of  vaccine 
was  sterile  after  one  hour  at  60°  C,  Possibly  the  animal  was  sensitized 
and  a  few  bacteria  settled  in  the  leg.  It  was  along  this  leg  that  the 
inoculations  were  made. 

We  permit  ourselves  the  facetious  observation  that 
that  vaccine  prevented  the  labial  and  cervical  variety  for 
five  years,  because  during  that  period  it  stood  in  the  ice- 
box, and  there  was  no  case  of  that  particular  form  to 
which  to  give  it,  although  a  few  of  the  nasal  and  gastric 
varieties  occurred.  It  was  recontrolled  and  did  not  show 
living  organisms.  That  it  should  have  cured  the  disease 
in  the  jaw  and  apparently  later  permitted  a  lighting  up 
of  a  septicemic  and  pulmonary  form  with  necroses  in  the 
leg  is  difficult  to  explain. 

Just  recently  we  have  used  a  vaccine  from  the  current 
culture  upon  another  case  beginning  in  the  gums  and  jaw 
bones.  This  case  was  detected  early  and  was  treated  with 
ascending  doses  beginning  at  0.3  mg.  and  running  up  to 
10.  mg.  At  first  there  was  some  improvement,  but  the 
animal  finally  died  from  pulmonary  complications.  The 
course  of  the  disease,  however,  instead  of  being  three 
weeks,  as  is  the  customary  duration,  lasted  two  months, 
an  extension  of  the  course  which  has  made  us  hopeful. 
These  two  experiments,  indefinite  though  they  be,  have 
offered  encouragement  and  seem  to  supply  a  little  addi- 
tional support  to  the  idea  that  the  organisms  stand  in 
etiological  relationship  to  the  disease. 

The  employment  of  the  vaccine  has  been  extended  to 
its  use  as  a  prophylactic  in  animals  exposed  to  the  disease 


594   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

or  specimens  that  have  slight  reddenings  or  erosions  on 
the  buccal  mucosae  suggesting  possible  early  stages  of 
streptothricosis.  Five  animals  have  now  had  a  course  of 
vaccine  injections,  ranging  in  number  from  5  to  10  and  in 
quantity  from  0.3  to  2.4  mg.  over  a  period  of  a  month. 
Fourteen  months  have  elapsed  at  the  time  of  writing  and 
only  one  case  has  developed,  but  this  of  course  cannot 
settle  the  efficacy  of  the  method ;  perhaps  it  would  be  safer 
to  demand  that  no  case  should  ever  appear  in  a  treated 
animal,  while  the  disease  did  appear  in  the  untreated. 

The  preparation  of  the  vaccine  is  by  no  means  a  simple 
matter,  since  the  surface  growth  upon  solid  media  is  so 
tenacious.  Methods  such  as  are  employed  for  the 
tubercle  bacillus  have  to  be  used.  The  first  two  vaccines 
were  made  by  scraping  off  surface  colonies  from  agar  and 
grinding  with  glass  balls.  One  successful  batch  was  made 
recently  by  simply  triturating  the  colony  directly  on  the 
agar  slant,  but  the  latest  method  seems  to  offer  the 
simplest  and  most  generally  satisfactory  way.  Neutral 
broth  is  placed  in  flasks  containing  glass  beads  and 
sterilized  in  the  incubator.  This  is  seeded  mth  the 
Nocardia,  incubated  at  37°  G.  until  the  surface  is  covered, 
heated  to  60°  C.  in  a  steam  sterilizer  and  tested  for 
sterility.  If  growth  occur  it  is  reheated  until  dead, 
whereupon  the  broth  is  syphoned  off,  the  growth  emulsi- 
fied by  whirling  the  flask,  thus  grinding  the  bacterial  mass 
by  the  glass  beads.  Sufficient  saline  is  added  to  make  a 
workable  emulsion,  and  the  fluid  then  poured  into  bottles. 
Control  by  reculturing  is  again  done,  and  if  the  fluid  be 
found  sterile,  0.5  per  cent,  trikresol  is  added  to  keep  it  so. 
These  organisms  cannot  be  counted  accurately  because 
of  the  variation  in  length,  their  budding  and  coccoid 
forms.  Standardization  is  done  by  weight.  A  definite 
equal  quantity  of  the  suspension  and  of  the  saline  used  to 
make  it  are  evaporated  to  dryness  in  weighed  vessels  and 
the  whole  then  weighed.  The  difference  is  the  weight  of 
the  organisms  suspended  in  the  saline.    Such  a  fluid  can 


THE  STREPTOTHRICOSES  595 

be  diluted  so  that  a  given  bulk  will  contain  a  convenient 
weight  of  germs.  The  one  now  in  use  contains  8.  mg. 
per  cubic  centimetre.  Dosage  as  indicated  above  usually 
begins  at  0.5  mg.,  a  quantity  which  does  not  produce  any 
local  inflammation  at  the  site  of  injection.  It  is  perhaps 
well  to  adopt  a  quantity  of  0.1  mg.  per  kilo  as  the 
initial  quantity. 

The  Garden  has  encountered  no  case  of  the  remaining 
important  chronic  infections,  glanders,  lymphangitis,  and 
infectious  abortion. 


SECTION  XVII— PART  4 

ACUTE  DISEASES  RESEMBLING  THE  SPECIFIC 
INFECTIONS  OF  DOMESTIC  ANIMALS 

Specific  communicable  diseases  are  sometimes  divided 
into  those  most  often  encountered  as  ''herd  diseases"  and 
those  which  appear  as  single  cases  or  in  small  groups. 
This  would  seem  to  imply  that  the  first  behave  as  easily 
disseminated  epizootics,  their  \drus  passing  from  animal 
to  animal  simply  by  proximity  or  by  casual  contact  where- 
as the  transfer  of  infective  material  is  less  readily  ac- 
complished by  the  second  group,  often  demanding  special 
assistance.  Foot-and-mouth  disease,  pleural  pneumonia, 
cattle  plague,  and  influenza  illustrate  the  epizootics  while 
tetanus,  rabies,  quarter-ill,  malignant  edema,  and  infec- 
tious vaginitis  are  examples  of  less  easily  transferred 
processes. 

It  is  not  intended  that  these  remarks  shall  cover  all 
possible  means  of  transmission  but  instead  they  are  in- 
tended to  focus  attention  upon  the  sources  of  viruses 
whereby  animals  become  infected.  An  original  case  must 
always  be  present  in  order  for  spread  to  occur.  Where 
animals  are  being  added  to  a  herd  a  new  comer  may  be 
diseased  or  the  carrier  of  a  virus;  when  animals  are 
transported  for  sale  or  other  reason,  infection  may  be 
met  in  a  new  stall,  conveyance  or  pasture ;  contaminated 
food  may  be  offered.  In  menageries,  with  specimens, 
single  or  in  small  groups,  and  arrivals  always  quarantined 
before  other  animals  are  exposed,  acute  specific  infections 
seldom  appear.  It  is  also  improbable  that  a  wild  animal, 
infected  at  its  source  or  in  some  dealer's  place,  would 
survive  the  journey  and  arrive  in  an  infective  condition. 
Consultation  with  the  reports  of  other  gardens  fails 
to  discover  records  of  any  serious  outbreaks  of  epizootic 
disease  except  for  fowl  cholera  and  distemper,  examples 

596 


MISCELLANEOUS  INFECTIONS  597 

of  infection  with  the  bipolar  organisms  of  the  Pasteurella 
group,  believed  responsible  for  the  hemorrhagic  septice- 
mias ;  instances  of  the  occurrence  of  the  group  specified 
secondly — anthrax  and  the  like — are  also  reported.  This 
represents  fairly  well  our  own  experience. 

The  bacteria  variously  named  Bac.  avisepticus,  ovisep- 
ticus,  hovisepticus,  canisepticus,  etc.,  grouped  by  Ligniere 
under  the  name  Pasteurella,  are  doubtless  of  considerable 
importance  and  are  probably  quite  widespread  in  natural 
surroundings.  The  viruses  of  the  epizootic  conditions 
like  cattle  plague  and  influenza  are  apparently  more 
definitely  parasitic,  requiring  for  their  persistence  ever 
renewed  transfer  from  host  to  host.  The  former  infec- 
tions we  have  met  in  repeated  single  isolated  cases  and 
in  small  groups,  whereas  no  cases  of  the  specific  epizootics 
have  been  diagiiosed. 

Hemorrhagic  septicemia,  a  denomination  very  descrip- 
tive of  its  pathological  picture,  has  been  encountered  in 
many  varieties,  carnivores,  ungulates,  primates,  rodents, 
and  birds.  The  diagnosis  depends  upon  the  presence  of 
hemorrhages  with  edema,  degenerations  of  the  parenchy- 
matous organs,  more  or  less  respiratory  catarrh  to  which 
may  be  added  relatively  mild  gastrointestinal  inflamma- 
tion; the  bacteria  are  found  in  the  circulating  blood  and 
in  exudates.  A  description  of  these  organisms  is  not 
profitable,  they  being  well  known  in  veterinary  pathology. 
"What  is  more  important,  significant  and  supportive  of 
the  opinion  expressed  above  concerning  the  mdespread 
distribution  of  the  virus,  is  the  incidence  of  the  infection. 
Exclusive  of  the  condition  known  as  fowl  cholera,  it  has 
appeared  among  mammals  and  birds  as  single  cases  with 
one  exception — that  of  two  Barbary  apes  which  had  been 
in  separate  cages  side  by  side.  The  total  of  cases  with 
determined  bacteriology  is  eleven,  with  undecided  bacte- 
riology but  suggestive  pathology  nine  additional.  No  per- 
tinent history  in  common  can  be  found  in  the  records 
of  the  determined  cases,  except  perhaps  that  they  were 


598  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

all  animals  which  had  been  in  the  collection  at  least  three 
months,  a  period  which  would  seem  to  exclude  the  proba- 
bility of  an  imported  infection.  Because  of  the  isolated 
character  of  the  cases  and  impossibility  of  making  a  clini- 
cal diagnosis,  no  attempt  at  specific  nomenclature  as  used 
in  veterinary  medicine  has  been  made,  hemorrhagic  sep- 
ticemia seeming  to  cover  its  identity  and  nature. 

The  disease  knowm  as  fowl  cholera  is  practically  al- 
ways associated  ^dth  the  bacteriological  discovery  of  a 
member  of  the  hemorrhagic  septicemia  group  while  its 
pathology  corresponds  with  that  of  mammalian  infection 
with  these  germs.  Enteritis  is  a  prominent  feature.  This 
disease  has  appeared  thrice  among  our  parrots  carrying 
off  from  six  to  ten  birds  before  hygienic  measures  became 
effective.  )  In  all  three  our  cultures  showed  the  bipolar 
organisms.  Besides  these  specific  outbreaks  numerous 
isolated  cases  of  acute  general  infection  have  occurred 
among  small  passerine  and  picarian  birds  which  could 
not  be  determined  as  hemorrhagic  septicemia  by  bacte- 
riological methods  although  superficially  resembling  it  in 
gross  pathology ;  they  yielded  to  the  same  hygienic  meas- 
ures. Perhaps  we  were  dealing  with  fowl  plague,  a 
disease  believed  to  be  due  to  a  filterable  virus.  That  this 
is  the  case  is  strongly  suggested  by  an  outbreak  of  fowl 
typhoid  in  the  parrots,  from  some  fatal  cases  of  which 
we  were  able  to  isolate  B.  sanguinarium,  and  by  a  group 
of  deaths  in  small  parrots  from  which  no  specific  organism 
could  be  recovered. 

The  identification  of  these  supposedly  specific  diseases 
— plague,  typhoid,  septicemia,  leucemia — by  pathological 
criteria  is  by  no  means  simple  even  if  we  have  at  hand  the 
complete  description  of  Moore,  of  Hutyra  and  Marek,  of 
EUermann  and  of  Ward  and  Gallagher,  Bacteriology 
must  decide  and  cultures  should  be  made  upon  bodies 
recently  dead.  In  addition  to  the  above  infections  we  have 
had  two  small  outbreaks  of  psittacosis  in  parrots  from 
which  it  was  possible  to  isolate  the  specific  organism.  On 


MISCELLANEOUS  INFECTIONS  599 

both  occasions  there  was  more  than  one  death  before  the 
specific  nature  of  the  disease  was  identified  yet,  note- 
worthily,  no  spread  to  the  other  birds  in  the  same  exhibi- 
tion house  occurred. 

Distemper,  a  disease  variously  held  as  due  to  cocci,  to 
influenza-like  organisms  and  to  a  filterable  virus,  may  ap- 
pear in  sporadic  or  epizootic  form.  The  diagnosis  during 
life  is  not  so  easy  unless  all  the  cardinal  features  are 
present,  while  after  death  the  same  thing  holds  good.  I 
am  inclined  to  think  that  from  the  standpoint  of  diagnos- 
tic accuracy,  the  term  is  used  much  too  loosely,  a  ready 
excuse  for  such  laxity  however  being  that  it  stimulates 
to  greater  care  in  hygiene.  Whether  or  not  B.  bronchi-  or 
canisepticus  be  the  cause  of  the  disease,  organisms  cor- 
responding to  it  can  be  found  in  stained  smears  from 
nearly  every  case  in  which  the  respiratory,  cutaneous, 
nervous  and  internal  signs  suggest  the  disease.  To  make 
a  diagnosis  of  distemper  it  is  my  practice  to  require  at 
least  three  of  the  cardinal  clinicopathological  features, 
whereupon,  if  the  bacterial  findings  be  as  described,  the 
denomination  is  permitted.  This  was  dictated  because 
during  the  period,  now  happily  well  in  the  past,  when  the 
cats  and  dogs  suffered  frequently  with  enteritis,  naso- 
pharyngeal signs  occasionally  presented  themselves  or 
spasms  were  reported,  but  no  skin  eruptions  appeared, 
yet  seldom  were  all  of  these  signs  combined  nor  could 
we  find  the  bipolar  organisms.  I  note  that  in  1915  Doctor 
Blair  of  New  York  observed  a  toxic  enteritis  resembling 
but  not  identical  with  distemper.  As  with  our  cases 
he  failed  to  find  that  the  condition  was  communicable. 
We  ascribed  our  cases  to  spoiled  food — fowl  heads  or 
dirty  horse  meat  (see  page  179).  Our  acceptable  examples 
of  distemper  number  three,  two  ferrets  and  a  lynx,  but 
very  suggestive  cases  were  found  in  foxes,  wolves  and 
raccoons.  Since  writing  the  above  notes,  sixteen  wolves, 
foxes  and  wild  dogs  died  in  an  outbreak  of  distemper 
imported  by  a  newly  arrived  specimen  admitted  to  the 


600   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

colony  by  mistake.  When  we  were  aware  that  the  disease 
had  appeared  antiserum  was  administered  therapeutically 
to  all  that  were  sick  and  prophylactically  to  all  the  rest 
— large  doses,  25-35  cc,  were  given  for  treatment,  smaller 
quantities,  10-20  cc,  being  used  as  a  preventive. 
Seven  sick  animals  recovered  and  no  animal  (8)  given 
serum  prophylactically  became  sick.  This  experience  en- 
courages us  to  think  that  with  antiserum  and  rigidly 
enforced  quarantine  rules,  distemper  will  not  be  a  serious 
matter  to  handle. 

The  hygiene  of  the  foregoing  conditions  is  of  a  general 
character — removal  of  the  specimens  when  kno^\^l  to  be 
sick,  thorough  cleansing  of  the  cages,  segregation  of 
mates  or  of  neighbors  when  this  is  practicable,  burning 
of  refuse,  liming  of  the  gromid  and  such  other  measures 
as  the  local  conditions  may  indicate. 

Diphtheria. 

Although  no  cases  of  mammalian  diphtheria  have  been 
observed,  three  and  possibly  four  birds  have  suffered 
with  tliis  disease.  The  three  acceptable  cases  were  in 
cassowaries  {Casuarius  occipitalis)  occupying  adjoining 
cages  and  sickening  within  a  few  weeks  of  one  another. 
Just  how  the  infection  was  brought  to  them  must  remain 
a  mystery  since  no  additions  had  been  made  to  the  group 
for  some  time  previously.  All  three  birds  were  observed 
during  life,  and  from  the  first  case  the  Bac.  diphtheria 
avium  was  isolated ;  in  smears  from  the  other  two  simi- 
lar bacteria  were  seen  but  isolation  was  unsuccessful.  The 
two  acutely  fatal  cases  showed  large  pseudomembran- 
ous collections  on  the  nasopharyngeal  mucosa  and  beneath 
the  tongue  while  the  nares  were  occluded  by  the  same 
material.  Plaques  of  membrane  were  also  found  on  the 
surface  of  the  esophagus  and  proventricle.  The  exudate 
ran  out  of  the  mouth  and  formed  dried  crusts  upon  the 
cervical  skin.  Pseudomembranes  of  a  continuous  charac- 
ter were  lacking  in  the  third  bird,  their  place  being  taken 


MISCELLANEOUS  INFECTIONS  601 

by  small  yellow  or  yellow-piuk  nodular  elevations,  appar- 
ently just  beneath  the  surface,  here  and  there  upon  the 
reddened,  slimy  buccal,  lingual  and  pharyngeal  mucosae. 
Crusts  upon  the  skin  of  the  neck  also  formed  in  this  case. ) 

These  cases  are  of  interest  not  only  because  of  their 
appearance  without  satisfactory  explanation  but  because 
one  improved  very  much  after  injections  of  human  diph- 
theria antitoxin,  this  remedy  being  used  because  we  were 
then  unaware  of  the  existence  of  an  avian  diphtheria  anti- 
toxin. No  claim  can  be  made  post  hoc  ergo  propter  hoc 
that  the  human  antitoxin  helped  the  attack — it  may  have 
been  mild — but  th^  experience  is  worth  recording.  Dosage 
was  as  follows:  '.December  3,  3,000  units;  December  8, 
1,500  units ;  December  21,  5,000  units ;  December  27,  5,000 
units.  Shortly  after  the  inception  of  the  treatment  the  bird 
was  noticed  to  eat  better  and  to  be  more  lively ;  this  was 
followed  by  a  reduction  in  the  mucous  strings  in  the  mouth 
and  the  crusts  upon  the  skin.  This  improvement  con- 
tinued and  the  bird  seemed  well  in  about  two  months  but, 
after  the  lapse  of  three  months  more,  a  mucous  naso- 
pharyngitis was  again  observed.  Despite  two  injections 
of  5,000  units  human  diphtheria  antitoxin  the  bird  suc- 
cumbed five  days  after  the  beginning  of  this  attack. 
Autopsy  revealed  much  the  same  condition  as  was  found 
in  the  first  birds  and  from  the  larynx  the  Bac.  avium  was 
isolated.  Another  case  suggestive  of  diphtheria  was  seen 
in  a  hornbill  but  antemortem  observation  being  imprac- 
ticable and  postmortem  decomposition  being  advanced 
when  autopsy  was  performed,  the  diagTiosis  could  not 
be  confirmed.  ;■ 

(An  unusually  well  developed  case  of  molluscum  con- 
tagiosum  was  seen  in  the  Wild  Turkey  (Meleagris  gallo- 
pavo)  recorded  here  by  photograph  and  in  the  form  of 
notes  upon  the  histology  made  by  Doctor  Weidman. 

The  bird's  head  was  affected  universally  from  beak 
to  ears  by  horny  nodules  up  to  the  size  of  a  pea.  They 
were  so  large  and  numerous  around  the  eyes  as  to  com- 

39 


602   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

pletely  close  them.  There  were  no  lesions  elsewhere  on 
the  body,  none  of  the  other  turkeys  were  similarly  affected 
and  though  watched,  none  have  since  developed  a  similar 
condition.  Histological  examination  shows  a  keratosis, 
many  of  the  cells  shomng  characteristic  *'  molluscum 
bodies  "  which  appear  the  same  and  behave  the  same 
tinctorially  as  the  human  examples.  Tliis  turkey  case 
differs  from  the  human,  however,  in  that  there  are  none 
of  the  pocket-like  epithelial  extensions  deep  down  into 
the  corium  and  this  turkey  case  may  be  very  useful  in  the 
further  study  wliich  is  contemplated  to  show  that  such 
things  as  molluscum  bodies  are  not  sufficient  of  them- 
selves to  stamp  a  dermatosis  as  a  pathological  entity,  but 
that  they  are  general  pathological  processes  wliich  may 
occur  in  a  number  of  different  diseases.  The  disease  has 
been  reported  in  sparrows,  pigeons,  but  never  so  far  as 
I  can  find,  in  turkeys.  ) 

A  few  isolated  cases  of  infectious  disease  are  included 
here  as  a  matter  of  record  although  they  may  not  be  espe- 
cially significant  or  important.  Rabies  was  found  in  a 
pair  of  deer  which  had  been  bitten  by  a  stray  dog.  The 
period  of  excitement  was  relatively  long,  while  the  para- 
lytic stage  was  only  a  few  hours.  Negri  bodies  were 
found.  Tetanus  killed  a  Persian  Wild  Ass  {Equus 
onager)  the  infection  wound  seeming  to  be  a  bruised  and 
abraded  area  on  the  rump.  From  the  contused  muscle 
tetanus  bacilli  were  isolated.  A  gas-bacillus  infection, 
emanating  from  the  vagina  which  was  protuberant  and 
lacerated  because  of  injury  by  mates,  was  seen  in  a  preg- 
nant llama  (Llama  lama.)  On  two  occasions  nodular 
masses  have  been  found  under  the  skin  of  seals,  not  unlike 
the  one  studied  by  Doctor  Wiedman  and  thought  by  him 
to  be  due  to  moulds.  These  two  have,  however,  failed  to 
show  mj^celia  or  yeast-like  bodies,  and  one  thinks  only  of 
placing  them  in  the  group  of  botryomycosis.  I  have  never 
seen  a  case  of  this  disease,  so  that  I  am  forced  to  rely  upon 
literature,  a  method  that  inspires  no  especial  confidence 


MISCELLANEOUS  INFECTIONS  603 

in  the  diagnosis.  The  bacteria  usually  held  responsible 
for  botryomycosis  could  not  be  isolated.  Just  what  can 
be  done  for  the  condition  is  difficult  to  state,  since  seals 
are  scarcely  tractable  animals. 

The  following  case  has  some  features  like  paralytic 
hemoglobinuric  fever  and  is  reported  as  a  matter  of 
record.  The  long  standing  gastroenteritis  may  have  been 
the  basis  for  the  intoxication  which  led  to  the  paralysis 
and  muscular  degeneration.  This  laboratory  has  now 
under  way  studies  upon  the  laming  of  ungulates,  accom- 
panied by  weakness  of  the  hind-quarters,  but  no  con- 
clusions have  been  reached.  It  is  interesting  to  note  that 
Hutyra  and  Marek  quote  Johne  as  having  seen  a  case  of 
hemoglobinuric  paralysis  in  a  zebra  in  1879. 

Burchell's  Zebra  {Equus  burchelli  burchelli).  The  only  symptom 
observed  in  this  animal  was  gradually  increasing  lassitude  which  was 
first  noticed  about  three  months  ago;  toward  the  end  he  habitually 
stood  with  tucked  tail  and  nose  to  the  ground  as  if  asleep.  He  ate  well 
and  digestion  appeared  good,  but  he  became  very  weak  as  shown  by  his 
inability  to  rise  when  he  got  down  on  the  third  and  second  day  before 
he  died,  although  on  both  occasions  he  was  able  to  stand  when  lifted. 
Injury,  hemorrhage  in  thigh  muscles,  chronic  gastritis,  sciatic  neuritis. 
CEstrus  larva  in  stomach,  ascaris  in  intestine.  Both  lungs  are  widely 
distended  and  the  caudal  half  of  both  is  the  seat  of  passive  congestion. 
Upper  lobes  are  slightly  edematous.  No  consolidations.  Heart  normal. 
Abdomen  contains  about  two  quarts  of  clear  straw  colored  fluid.  Liver 
is  of  normal  size,  smooth  surface,  sharp  edges,  firm,  friable.  On  section 
it  is  very  bloody,  veins  distended,  some  with  clot.  Architecture  normal. 
Spleen  is  of  normal  size,  soft,  tough,  capsule  rough.  Section  surface  is 
homogeneous,  pulp  purple,  trabeculae  normal,  follicles  not  visible.  The 
kidney  capsule  is  smooth,  strips  easily  leaving  a  smooth  brown  surface, 
firm.  StriiE  nonnal,  rather  wide,  glomeruli  not  visible.  Stomach  is 
(filled  but  not  distended  -with  partly  digested  straw.  Mucosa  of  cardia 
dry,  roughly  irregular,  some  irregular  mammillations.  Two  flat  papil- 
lary growths.  (Estrus  larva  attached  to  a  smaller  elevation.  The 
mucous  membrane  of  the  fundus  is  soft,  moist,  irregular,  in  some  places, 
translucent,  in  others  opaque ;  near  pylorus  mucous  membrane  is  swollen 
edematous,  pink,  slightly  eroded  at  pyloric  valve.  Small  intestine  has 
smooth,  flat,  pale  yellow  translucent  mucosa.  Lumen  filled  with  muco- 
purulent matter  like  mixed  egg.  Ileum  slightly  congested  but  mucosa 
firm  and  translucent.  Pancreas  is  soft,  slightly  uniformly  congested. 
All  mesenteric  lymph  glands  are  slightly  enlarged  and  edematous  but 
with  normal  architecture.     In  the  posterior  thigh  muscles  beside  the 


604   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

sciatic  nerve,  most  marked  on  the  right  side,  is  a  large  hemorrhagic 
infiltration.  There  is  edema  of  muscles  and  intermuscular  septa  all 
about  this  area  extending  upward  as  Avell  as  to  pelvis  and  psoas  muscle. 
This  latter  within  the  abdomen  shows  slight  blood  stained  edema.  No 
other  muscle  shows  this  hemorrhage.  Microscopic  section  of  liver  and 
kidney  are  negative  aside  from  congestion.  The  stomach  shows  very 
irregular  epithelial  covering,  in  some  places  wholly  desquamated.  Where 
this  is  most  marked  there  is  a  dense  round  cell  infiltration  in  the  villi 
with  some  increase  in  the  connective  tissue  cells.  This  chronic  inflamma- 
tory reaction  is  present  in  all  fields,  most  marked,  of  course,  in  upper 
layers  of  mucosa.  Glands  are  distorted  and  upper  epithelium  of  them 
is  polychromatophilic.  The  intestine  shows  similar  changes  in  less  in- 
tense manner. 

Waterfowl  Epizootic.  There  is  reproduced  here  an 
account  of  an  unexplained  epizootic  among  ducks  and 
geese  from  the  Annual  Report  of  the  Zoological  Society 
for  1916.  Nothing  additional  has  been  learned  and  no 
repetition  has  occurred  since  the  drainage  and  cleaning 
of  the  lake. 

There  began  on  August  27  a  series  of  deaths  among  the  waterfowl 
and  in  one  month  there  were  lost  forty-one  specimens  including  both 
ducks  and  geese.  Four  additional  cases  were  scattered  through  the  next 
four  months,  the  last  case  dying  January  11,  1916.  All  of  these  came 
from  the  lake,  none  being  from  the  adjacent  stream  for  rare  waterfowl 
or  from  the  more  distant  stream  into  Avhich  the  lake  drains.  The  sj^mp- 
toras  were  most  marked  and  striking.  In  the  early  stages  the  wings 
drooped,  then  the  legs  became  weak  followed  by  inability  to  raise  the 
head.  In  the  latest  cases  the  voice  (ducks)  lost  its  normal  character  and 
became  hissing.  The  mind  appeared  clear  for  the  eyes  were  bright, 
feathers  unruffied  and  the  bird  attempted  to  escape  when  approached. 
Diarrhoea  was  present,  dejecta  thin,  watery  white,  no  admixture  of 
mucus.  Autopsy  findings  were  not  frank.  At  most  some  swelling  of  the 
spleen  and  a  little  pale  thickening  of  the  intestinal  wall  constituted  the 
picture.  Smears  from  intestine  and  nasal  mucosa  showed  no  protozoa. 
The  blood  taken  from  the  living  sick  ducks  showed  no  parasites  or  anemic 
changes  in  either  raw  or  variously  stained  preparations.  From  the 
spinal  cords  of  three  ducks  a  50  per  cent,  glycerine  emulsion  Avas  pre- 
pared and  Avas  injected  into  the  cerebral  substance  and  abdomen  of 
domestic  ducks  with  negative  results.  A  variety  of  different  bacterial 
cultures  was  obtained  from  the  liver,  spleen,  blood  and  congested  nasal 
mucosa  of  several  birds  dead  with  the  disease  and  injected  into  domestic 
ducks  with  negative  results.  Histological  sections  were  cut  from  the 
important  organs  of  thirteen  birds.  The  kidneys,  lungs  and  pancreas 
shoAved  no  abnormalities.     The  heart  muscle  in  some  cases  and  also 


MISCELLANEOUS  INFECTIONS  605 

some  of  the  skeletal  muscles  showed  Zenker's  hyaline  degeneration 
together  with  minor  hemorrhages  and  edema.  Several  of  the  proven- 
trieles  showed  low  grade  inflammatory  signs  toward  the  gizzard.  The 
intestines  regularly  showed  lymphatic  infiltrations  of  the  villi  most 
marked  toward  the  tips  but  without  congestion.  The  lumen  showed  no 
parasites,  bacteria  or  protozoa.  Liver  showed  in  almost  every  case 
pigmentation  by  hemosiderin  at  times  as  heavy  as  that  seen  in  per- 
nicious anemia.  The  finer  bile  ducts  here  showed  peripheral  round  cell 
infiltrate,  which  was  not  continued  into  the  major  ducts  as  determined 
by  serial  sections.  Parenchymal  cells  were  cloudy  and  swollen.  Spleen 
showed  in  early  cases  polymorphonuclear  infiltrate  of  the  follicles,  in 
later  cases  atrophy  of  follicular  splenoeytes  and  more  or  less  pigment 
occun-ed  in  both  stages.  The  spinal  cord  and  various  peripheral  nerves 
showed  no  inflammation  or  degeneration  as  determined  by  the  appro- 
priate special  nerve  stains.  The  above  clinical,  histological,  protozoologi- 
cal,  and  bacteriological  examinations  having  failed  to  detect  the  cause 
and  the  epizootic  now  being  over,  its  nature  becomes  a  matter  of 
deduction.  The  only  constant  features  of  any  importance  were  the 
paralysis,  the  intestinal  round  cell  infiltrate  and  thickening,  the  pigmen- 
tation of  the  liver  and  degeneration  of  skeletal  muscles.  Of  the  various 
possibilities,  beriberi  was  early  considered.  This  is  not  possible  because 
the  food  of  the  birds  was  a  varied  one  and  furthermore  none  of  the 
neiwe  degenerations  of  beriberi  were  noted.  Second,  acute  bacterial  or 
protozoal  infections  are  unlikely  because  no  constant  primary  lesions 
were  discovered  at  autopsy,  the  numerous  cultures  failed  to  produce  the 
disease  and  other  birds  living  on  the  stream  draining  the  lake  were  not 
similarly  affected.  Third,  a  food  poisoning.  This  is  possible  first  because 
paralytic  symptoms  were  present  such  as  are  seen  in  vetch  and  mussel- 
poisoning  and  secondly  because  the  epizootic  ceased  when  the  birds  were 
taken  from  the  lake  and  placed  upon  the  grass.  If  this  be  the  case  the 
toxic  material  produced  the  paralysis  by  direct  action  upon  the  muscle 
fibres  just  as  that  of  typhoid  fever  does  and  must  have  caused  hemolysis 
as  shown  by  the  hepatic  pigmentation.  The  source  of  this  food  poison- 
ing is  conjectural.  Perhaps  a  dead  fish  decomposed  in  the  water  or 
there  were  some  algae  with  poisonous  properties  present.  The  outbi-eak 
has  a  resemblance,  but  only  a  superficial  one,  to  infection  with  one  of 
the  group  of  botulism  bacilli.  The  cause  of  the  trouble  must  be  con- 
sidered as  undetermined. 

Enterohepatic  Disease.  Since  the  normal  drainage 
from  the  intestinal  tract  passes  so  largely  through  the 
liver,  there  is  little  to  wonder  at  in  morbid  lesions  of  the 
latter  organ  consequent  upon  disease  in  the  former.  Not 
only  does  this  succeed  upon  bacterial  infection  of  the 
digestive  tube  but  also  upon  infestation  with  animal 
parasites,  under  the  latter  condition  forming  changes  of 


606   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

much  more  considerable  extent,  at  least  in  gross  bulk, 
than  in  the  former.  Changes  in  the  liver  secondary  to 
enteric  disease  from  bacterial  infection  take  the  form  of 
cholangitis,  thrombosis,  degenerations  and  probably 
cirrhosis  while  abscesses  and  necroses  succeed  upon  pro- 
tozoal or  metazoal  parasitic  involvement.  JThe  latter  is 
exemplified  by  amebic  abscess  in  man  and  other  mam- 
mals and  by  ''blackhead"  and  ''quail  disease"  in  birds; 
it  is  to  the  latter  conditions  that  attention  is  now  directed. 
The  chapter  upon  the  cause  of  these  diseases  has  yet  to 
be  completed,  although  many  reams  have  been  written 
about  it,  while  the  transmission  is  fairly  well  understood 
and  the  pathology  well  described.  My  purpose  here  is  to 
discuss  our  experience  with  the  two  above  mentioned 
diseases  which,  while  far  from  conclusive,  may  assist 
somewhat  in  explaining  their  etiology.  There  is  also 
reproduced  our  original  report  upon  quail  disease  from 
the  Society's  Report  of  1915,  giving  data  and  figures. 
(  Blacldiead  has  been  found  in  five  wild  turkeys.  An 
unusual  case  in  a  Berwick's  Swan  is  recorded  since  it 
bears  a  striking  resemblance  to  the  disease. 

The  points  at  issue  in  the  determination  of  the  etiology 
of  blackhead  are  the  importance  of  Heterakis  papillosa 
in  the  ceca  and  the  frequency  and  activity  of  ameba  or 
histomonas. '  In  three  of  the  five  cases  of  the  disease  in 
turkeys  the  nematode  was  found  macroscopically  in 
the  ceca,  in  two  it  was  not;  in  one  its  absence 
was  confirmed  microscopically.  In  two  of  the  turkey 
cases,  forms  corresponding  to  the  ameba  or  histomonas 
were  discovered  while  the  descriptions  of  the  he- 
patic lesions  in  two  birds  use  the  term  coccidia  which, 
from  a  revision  of  the  slides,  is  probably  incorrect 
although  some  of  the  parasites  seem  to  be  possessed  of  a 
doubly  contoured  refractile  margin.  The  larger,  more 
diffuse  and  ameba-like  forms  in  the  intestinal  wall  sug- 
gest that  the  hepatic  inclusions  belong  to  the  same  group. 
In  only  one  case  was  exhaustive  search  made  for  coccidia, 


MISCELLANEOUS  INFECTIONS  607 

and  without  success ;  the  material  was  not  preserved.  In 
two  turkeys  entirely  free  of  lesions  distinctive  of  black- 
head, cecal  nematodes  (one  heterakis,  one  unknown)  are 
recorded,  and  in  the  intestinal  wall  of  another,  also  free 
from  the  disease,  forms  indistinguishable  from  ameba 
could  be  discovered.  (1) 

The  protocol  of  the  Berwick's  Swan  is  interesting 
because  the  full  fledged  disease  is  not  known  in  this  bird. 
While  this  case  is  not  by  any  means  typical,  the  chronic 
cecitis  and  ameba-bearing  necroses  in  the  liver  stamp  it 
as  of  a  kind  with  the  true  infection  of  turkeys.  Perhaps 
the  resistance  offered  by  the  swan  effected  a  modification 
of  the  disease,  preventing  the  usual  necrotizing  enteritis 
and  turning  it  into  a  chronic  interstitial  variety. 

Berwick's  Swan  {Cygnus  herwicki).  About  a  month  before  death 
passed  several  large  clots  of  blood.  Acute  catarrhal  enteritis,  mural 
endocarditis,  chronic  colitis,  chronic  nephritis,  passive  congestion  and 
necroses  in  liver,  acute  follicular  splenitis,  edema  of  lungs,  chronic  peri- 
carditis, chronic  salpingitis,  hydrothorax,  hydropericardium,  hydro- 
peritoneum.  Tissues  generally  are  slightly  yellow.  In  serous  cavities 
of  thorax  is  about  three  ounces  of  clear  fluid.  Lungs  are  distended, 
subcrepitant,  pale  red  and  gray,  highly  edematous.  The  pericardium 
contains  about  one-half  ounce  of  clear  watery  fluid.  Epicardium  is 
glistening,  congested,  irregularly  thickened  especially  near  the  blood 
vessels.  The  heart  is  contracted,  slightly  large,  pale  brown-red  muscle. 
On  the  posterior  surface  of  the  right  ventricle  extending  from  the  auric- 
ular opening  to  the  pulmonaiy  valve  is  an  irregularly  curved  line  of 
grouped,  recent  red  vegetative  granulations.  Valves  negative,  they  and 
chambers  competent.  Aorta  negative  except  heavily  blood  stained. 
Liver  is  slightly  large.  What  of  the  liver  remains  undamaged  is  homo- 
geneous deep  purple.  Major  portion  of  right  lobe  badly  contused ;  this 
seems  to  have  been  partly  antemortem  because  there  is  blood  staining 
and  mottling  under  capsule.  In  view  of  colon  finding  and  history  of 
possible  injury  it  is  probably  the  result  of  degenerations  in  the  liver  plus 
slight  trauma.  There  are  several  small,  pale  gray,  well  outlined,  homo- 
geneous areas  probably  necroses  in  the  liver.  The  spleen  is  slightly 
large,  soft,  egg-shape,  capsule  smooth.  Section  surface  shows  bright 
red  homogeneous  pulp  with  clearly  cut,  large  follicles.  The  kidney 
capsule  is  smooth,  surface  smooth  brown,  consistency  firm  and  tough. 
The  section  surface  gives  a  dull  gray-brown  appearance,  seemingly  from 

( 1 )  Those  interested  in  the  investigation  of  the  cause  of  Blackhead 
are  referred  to  the  recent  literature  by  Tyzzer  and  by  Smith,  in  the  Jour, 
of  Exp.  Med.  and  Jour,  of  Med.  Research,  1918-1922. 


608   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

connective  tissue.  Markings  indistinct.  Oviduct  is  negative  except  over 
a  distance  of  an  inch  near  the  cloacal  opening.  Here  there  is  a  com- 
pound curve  with  constriction  to  almost  obliteration  of  lumen.  This 
does  not  seem  to  be  connected  with  the  colonic  trouble.  The  stomach  is 
negative  containing  only  a  few  small  pebbles.  Beginning  at  the  pylorus 
and  extending  through  the  whole  of  the  small  gut  is  a  recent,  moderately 
severe  catarrhal  enteritis  with  so  much  exudate  as  to  form  almost  a  cast 
of  the  tube.  Colon  and  cloaca  show  an  infiltration  of  submucosa  with 
areas  of  hemorrhage.  Mucosa  swollen  as  if  by  edema,  glistening  and 
covered  by  bloody  mucus.  Ceca  negative  except  that  they  seem  to  have 
been  closed  as  their  contents  are  scanty  and  firm.  Histological  section 
of  cloaca  shows  it  to  be  the  seat  of  a  chronic  inflammation  which  has 
constricted  and  distorted  the  tubules  into  simple  masses  of  nuclei. 
Marked  polynuclear  and  round  cell  infiltration  of  mucosa  and  submucosa. 
This  is  apparently  due  to  ameba-like  bodies — a  large  vacuole  with  a 
delicate  limiting  membrane  and  a  piece  of  diffuse  chromatin  in  the 
centre — a  few  of  which  may  be  found  deep  in  the  mucosa.  Liver  shows 
marked  passive  congestion,  here  and  there  areas  of  necrosis  with  some 
fatty  infiltration.  Small  groups  of  ameba-like  bodies  can  be  found 
apparently  lying  in  sinusoids  of  liver  and  in  neighborhood  of  necroses. 

Quail  disease,  since  the  careful  work  of  Morse  in  1907, 
has  been  thought  by  most  observers  to  be  due  to  an  organ- 
ism of  the  colon  group,  but  I  am  informed  recently  by  the 
Pennsylvania  State  Board  of  Animal  Industry  that 
coccidia  have  been  found  often  enough  in  the  droppings 
and  in  the  morbid  lesions  to  warrant  a  suspicion  of  their 
etiological  importance.  Although  they  were  not  espe- 
cially sought  in  the  work  about  to  be  reported,  their  pres- 
ence probably  would  not  have  escaped  detection  during 
that  investigation.  I  have  recently  had  occasion  to 
examine  three  birds  with  lesions  identical  with  those 
accepted  as  characteristic  of  quail  disease,  one  of  which 
was  subjected  to  the  proverbial  ''fine  tooth  comb" 
methods ;  no  coccidia  were  found  in  the  liver  or  intesti- 
nal lesions. 

The  idea  that  quail  disease,  with  its  ulcerative  typh- 
litis and  necrotizing  hepatitis,  is  identical  with  blackhead 
or  at  least  that  if  the  latter  be  due  to  protozoa,  the  former 
is  also,  requires  no  special  stretch  of  imagination  to  one 
famiUar  with  the  morbid  lesions.  A  decision  is  the  more 
difficult  because   of  one's  inability  to  reproduce  quail 


MISCELLANEOUS  INFECTIONS  609 

disease  at  will  and  the  none  too  great  certainty  of  the 
intentional  production  of  blackhead.  At  all  events  the 
transmission  is  potentially  the  same,  gromid  or  food 
soiled  with  droppings,  indicating  that  hygienic  measures 
should  take  the  form  of  segregation  and  disinfection. 
Here  follows  the  report  of  our  original  observation : 

'^  An  epizootic  disease  has  decimated  three  newly  im- 
ported lots  of  quail,  Scaled  quail  (Callipepla  squamata), 
GambePs  quail  {Lophortyx  gamheli)  and  Texas  bobwhite 
{Colinus  texasus  virgimanus).  On  January  5,  1915, 
the  first  lot  of  twenty-four  quail  arrived  from  northern 
New  Mexico  via  Kansas  City;  on  Januaryllth  a  second  lot 
of  twelve  bobwhite  arrived  from  Brownsville,  Texas,  via 
Kansas  City ;  the  first  of  this  lot  died  the  day  after  arrival 
with  lesions  of  this  infection.  From  this  lot  of  birds  the 
first  lot  was  probably  infected,  the  first  death  occurring 
on  January  20th,  no  other  deaths  having  occurred  in  the 
first  lot  since  arrival.  On  January  21st  the  third  lot  of 
twelve  quail  arrived  direct  from  Mexico.  The  first  of  this 
lot  died  of  the  disease  on  January  24th.  Some  birds  were 
also  sent  at  the  time  of  the  arrival  of  the  third  consign- 
ment, to  Doctor  Kalbfus  of  the  State  Game  Commission. 
It  is  to  be  emphasized  that  to  date  no  cases  of  infectious 
enteritis  have  occurred  in  the  lot  sent  to  Doctor  Kalbfus. 
The  first  case  appeared  at  this  Garden  on  January  12th, 
more  than  a  week  before  the  third  lot  arrived.  It  would 
seem  that  the  disease  was  brought  to  the  Garden  by  the 
second  lot  of  birds,  and  that  they  picked  it  up  on  the  way 
from  Texas  to  Kansas  City  to  Philadelphia.  The  birds 
made  a  stop  at  Kansas  City.  The  birds  died  at  long 
intervals  for  the  first  two  weeks,  but  late  in  January  and 
early  in  February  several  died  each  day.  The  last  death 
with  characteristic  lesions  occurred  February  11th.  After 
the  epidemic  reached  its  height  it  subsided  very  quickly. 

''During  the  illness  the  birds  exhibited  very  few  symp- 
toms, indeed  some  of  them  were  not  known  to  be  sick.  A 
few  sat  huddled  in  a  corner  mth  ruffled  feathers  and 


610  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

drooping  head;  the  stools  were  little  if  any  altered  as  far 
as  could  be  determined  among  so  many  in  the  enclosure. 
At  death  the  birds  were  in  good  condition,  feathers  fairly 
smooth,  skin  clear,  body  plump  and  fat  in  good  amount — 
not  abundant,  nor  were  the  animals  emaciated.  The 
principal  lesions  were  enteritis,  degenerative  necroses 
and  abscesses  in  the  liver,  congestion  of  all  the  viscera 
and  plastic  peritonitis  in  a  few.  A  small  number  showed 
congestion  of  the  Imigs  and  two  had  patches  of  pneu- 
monia. Many  but  not  all  of  the  birds  had  Heterakis  in  the 
ceca.  The  process  seemed  to  start  as  a  focal  necrotizing 
lesion  in  the  mucosa  or  submucosa  of  the  ileum  just  above 
the  ceca  and  colon ;  many  had  lesions  in  the  ceca  and  as 
far  down  in  the  colon  as  the  cloacal  dilatation.  Among  the 
animals  dying  late  in  the  epidemic  several  showed  lesions 
involving  the  whole  small  intestine,  a  few  indeed  with 
greater  involvement  of  the  duodenum  than  of  the 
lower  parts. 

(** Judging  from  the  gross  and  microscopical  appear- 
ances it  seems  that  the  virus  causes  at  first  a  cellular  infil- 
trate in  the  mucosa  or  submucosa  upon  which  necrosis 
shortly  supervenes.  The  overlying  mucosa  soon  degen- 
erates, and  the  surface  is  covered  with  an  indefinite 
slough.  In  other  cases,  especially  early  in  the  epidemic, 
the  process  extended  outward  and  appeared  as  muscular 
or  subperitoneal  necrotic  areas  before  the  mucosa  was 
much  involved.  At  all  events  necrosis  was  an  early 
change  in  every  case.  The  blood  vessels  were  usually 
thrombotic.  In  the  cases  that  spread  toward  the  peri- 
toneum a  plastic  peritonitis  of  varying  severity  was 
present.  The  focal  liver  lesions  were  not  present  in  every 
case.  They  took  the  form  of  focal  necroses  or  abscesses. 
Some  fatty  or  parenchymatous  degeneration  w^as  always 
present.  The  liver  lesions  probably  started  as  inflamma- 
tions of  the  veins  from  which  necrotizing  or  infiltrative 
lesions  spread.  The  splenic  lesions  were  those  of 
lymphoid  hyperplasia,  only  distinctive  in  the  enormous 


MISCELLANEOUS  INFECTIONS  611 

number  of  large  lymph  cells.  Typical  microscopical 
changes  are  as  follows,  quoted  from  one  of  the  autopsy 
protocols :  The  lung  showed  moderate  congestion  with 
here  and  there  a  little  epithelial  swelling  and  a  mild  bron- 
chitis and  peribronchitis.  The  type  of  bronchitis  is 
infiltrative  rather  than  catarrhal.  The  heart  muscle 
showed  granular  degeneration  of  the  fibres  with  breaking 
up  or  irregularity  of  the  striae.  Some  increase  in  inter- 
fibrillar  nuclei  and  especially  those  of  the  capillaries. 
There  is  moderate  congestion.  Epi-  and  endocardia  are 
slightly  raised  as  if  by  edema.  Here  and  there  slight 
fragmentation  of  fibres.  The  liver  cells  are  granular 
and  some  show  fat  droplets.  There  is  moderate  conges- 
tion and  more  than  the  normal  number  of  round  nuclei 
between  the  columns.  Here  and  there  are  focal  necroses 
of  varying  sizes  without  circumferential  reaction.  Here 
and  there  are  also  some  small  collections  of  round  cells 
near  to  which  the  liver  nuclei  are  large  and  show  attempts 
at  regeneration.  In  these  collections  but  not  in  the 
necroses,  bacillary  forms  may  be  found.  There  is  no 
reaction  on  the  part  of  the  bile  ducts.  The  larger  vessels 
are  thrombotic,  and  in  one  section  a  thromboangiitis  was 
found.  One  stretch  of  early  plastic  perihepatitis  was 
found.  The  kidney  showed  slight  granularity  with  slight 
cloudy  swelling  of  the  epithelium.  The  nuclei  of  the 
glomeruli  are  prominent.  There  is  moderate  congestion. 
The  spleen  showed  distinct  large  lymph  cell  hyperplasia 
with  relative  inconspicuousness  of  small  round  cells. 
The  follicles  are  very  diffuse,  their  centres  filled  with 
large  lymph  cells.  The  cords  are  hyperplastic  and  the 
sinuses  compressed.  Moderate  congestion;  no  unusual 
blood  destruction;  one  area  of  hyaline  necroses  found. 
The  proventricle  and  gizzard  are  negative  with  the  prob- 
able exception  of  active  desquamation  on  the  surface  of 
the  former.  The  outer  coats  of  the  duodenum  are  nega- 
tive except  for  slight  richness  in  nuclei.  The  deep  mucosa 
is  very  rich  in  nuclei  and  red  blood  cells.    The  outer  parts 


612   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

of  the  villi  are  either  swollen  with  a  cellular  infiltrate  or 
by  an  area  of  granular  necrosis,  or  have  disappeared.  It 
would  seem  that  the  surface  of  the  mucosa  rapidly  degen- 
erates and  desquamates.  Bacteria  are  very  numerous. 
The  adjacent  pancreas  is  negative.  The  ileum  showed 
round  cell  infiltration  of  the  deep  mucosa,  swelling  of  the 
villi  and  a  desquamation  of  the  surface.  One  ulcer  was 
found  having  its  base  on  the  swollen  muscularis  and 
being  covered  with  necrotic  slough.  Adjacent  peritoneum 
is  slightly  infiltrated,  but  chiefly  congested  and  edema- 
tous. This  ileum  lesion  seems  to  be  the  characteristic  one 
of  the  disease.  Bacteriological  observations  were  made 
upon  cultures  obtained  from  the  intestinal  mural  lesions, 
the  peritoneal  exudate,  the  liver  necroses,  and  the  heart's 
blood  in  eleven  cases.  In  seven  cases  I  was  able  to  isolate 
a  motile  rod  like  the  B.  coli  communis  and  in  four  cases 
a  non-motile  rod  of  the  Bad.  aero  genes  type.  The 
former  is  quite  similar  to  the  B.  scoticus  (Migula) 
reported  in  Grouse  disease. 

*'We  obtained  from  Doctor  Kalbfus  of  the  Pennsyl- 
vania State  Game  Commission,  four  perfectly  healthy 
birds  for  experimentation.  A  culture  of  the  isolated  germ 
was  injected  into  two  of  them  and  mixed  with  the  food  of 
the  remaining  two.  It  does  not  seem  profitable  to  cite  the 
details  of  the  work  as  the  results  were  entirely  negative, 
no  lesions  resulting  that  bore  the  slightest  resemblance  to 
the  spontaneous  disease.  .  The  birds  either  lived  indefi- 
nitely or  succumbed  to  wholly  foreign  conditions.  This 
negative  experiment  is  of  course  no  proof  that  the  organ- 
ism is  not  the  cause  of  quail  disease,  for  the  methods 
employed  might  not  be  the  correct  ones  to  propagate  the 
virus  or  the  germ  may  have  lost  its  virulence  during  the 
laboratory  culture  work.  However,  as  some  observers 
have  not  reported  this  bacillus  in  the  disease  this  germ 
loses  something  in  importance  by  the  negative  inocula- 
tion experiment. 


MISCELLANEOUS  INFECTIONS  613 

"Judging  from  reports  and  based  upon  the  observa- 
tions of  Morse  upon  Grouse  disease  it  would  seem  that  the 
incubation  period  of  the  disease  is  about  eight  to  ten 
days.  However,  one  of  the  tliird  lot  of  our  birds  died 
within  three  days  of  its  arrival  at  this  Garden,  and  there- 
fore within  three  days  of  its  exposure  to  the  second 
arrivals ;  if  it  be.  correct  that  this  second  lot  brought  the 
disease  and  the  third  lot  did  not  have  it,  it  would  seem 
that  the  incubation  period  can  be  as  short  as  three  days ; 
how  long  it  may  be  is  only  suggested  by  the  fact  that  some 
of  the  third  lot  did  not  die  for  three  weeks  after  arrival 
and  exposure.  All  the  Gambel's  and  scaled  quail  suc- 
cumbed to  the  disease,  but  two  of  the  twelve  bobwhite 
survived.  It  would  seem  that  although  these  last  birds 
probably  introduced  the  disease,  they  still  possessed  more 
resistance  than  the  others,  for  the  second  death  among 
them  occurred  seventeen  days  after  the  first  death./;  The 
epidemic  as  we  have  seen  it  here  seems  to  be  the  same 
as  Grouse  disease  of  Scotland  and  as  the  Grouse  disease 
in  this  country  as  reported  by  Morse  (Bureau  of  Animal 
Industry  Report  109,  May  18,  1907). 

"The  means  of  transmission  of  the  disease  is  not 
exactly  known,  but  is  in  all  probability  by  a  pollution  of 
the  food,  the  water  supply  or  the  ground.  Since  the  lesions 
are  so  marked  in  the  lower  ileum,  cecum  and  colon,  a 
possible  transmission  by  cohabitation  must  not  be  entirely 
overlooked.  There  does  not  seem  to  be  any  means  of 
limiting  the  epidemic  in  a  flock  by  segregation  or  sacrifice 
of  the  infected  birds,  because  symptoms  are  few  and  do 
not  appear  until  shortly  before  death.^  Each  bird  would 
have  to  be  put  into  a  separate  cage  until  proved  infected. 
Scrupulous  cleansing  of  the  enclosure  is  desirable,  but  its 
efficiency  is  difficult  to  estimate." 


SECTION  XVIIl 

THE  ANIMAL  PARASITES,  THEIR  INCIDENCE 
AND  SIGNIFICANCE 

Feed  D.  Weidman,  M.  D. 

It  is  quite  to  be  expected  that  animal  parasites  would 
be  found  in  the  animals  of  zoological  gardens,  garnered 
as  these  beasts  are  from  all  parts  of  the  world,  tropical 
and  otherwise.  It  inevitably  follows  that  many  of  the 
forms  should  be  strange  and  new,  enticing  one  to  the 
fascinating  determination  of  their  identity,  life  history 
and  hygienic  importance ;  and,  developing  from  all  this, 
one  can  easily  imagine  how  limitless  the  opportunities 
are  for  scientific  work  in  parasitology  in  a  laboratory 
like  ours. 

As  in  other  biological  fields,  the  taxonomic  range  of 
parasites  here  is  vvide.  It  extends  fr'om  the  lowly  proto- 
zoa to  the  insecta,  and,  dropping  to  the  smaller  subdivi- 
sions, includes  not  only  most  of  the  genera  familiar  to 
human  parasitology  but  many  knov\Ti  only  among  the 
lower  animals.  From  the  standpoint  of  the  host,  the  bio- 
logic state  of  parasitism  extends  from  the  lowest  protozoa 
to  homo. 

The  above  vnll  suffice  to  indicate  the  wide  range  of 
parasitism  in  animals,  but  the  extent  of  work  actually 
done  thus  far  in  wild  animal  material  is  a  different  story. 
Collated,  consistent  studies,  so  far  as  I  am  aware,  have 
been  undertaken  only  at  the  London  Garden,  here  at 
Philadelphia,  and  at  Wasliington,  D.  C,  by  Dr.  Charles 
W.  Stiles  and  Albert  Hassal.  The  data  collected  by  the 
last  mentioned  workers  are  incidental  to  the  Index  Cata- 
logue of  Veterinary  and  Medical  Zoology,  and  embrace 
only  the  (index)  phase  indicated  by  the  title,  but  it  is  so 
valuable,  and  vvithal  so  altruistic,  that  it  must  be  credited. 

614 


THE  ANIMAL  PARASITES  615 

What  other  work  there  is  is  scattered  where-not  in  lit- 
erature— general  biological,  medical  and  veterinary.  That 
at  London  has  been  conspicuous  through  the  observations 
of  Plimmer  and  of  Beddard  on  filariae  and  cestodes 
respectively,  while  the  w^ork  of  Nicoll  must  not  fail 
of  mention. 

That  the  reader  may  the  better  appraise  the 
sections  of  our  own  work  which  are  to  follow  I  wish 
at  once  to  indicate  their  material  basis.  Ordinarily 
only  the  larger  parasites  are  looked  for  at  the  autopsy 
table  and  there  must  be  special  indications  to  demand 
search  for  the  finer  ones.  Those  of  microscopic  size, 
or  so  minute  as  to  be  overlooked  in  the  guise  of 
seeds,  vegetable  fibres,  etc.,  have  not,  both  here  and 
elsewhere,  been  routinely  studied  as  have  macroscopic 
ones.(l)  From  our  autopsies  there  have  accumulated 
records  of  nearly  900  parasites — some  determined  generi- 
cally,  others  but  as  to  order.  The  parasites  have  in 
greatest  part  been  preserved  and  are  available  for  fur- 
ther study ;  in  the  past,  special  groups  have  been  culled 
out  from  time  to  time  and  examined.  Where  conditions 
have  been  pressing,  as  in  certain  epizootics,  investigations 
have  amounted  to  more  than  observations  and  descrip- 
tions, and  received  detailed  laboratory  examinations  with 
more  or  less  animal  experimentation  as  the  occasion 
demanded. 

The  foregoing  may  suffice  to  apprise  the  reader  that 
the  subject  of  wild-animal  parasites  has  been  but  broached 
so  that  data  are  especially  incomplete  on  life  histories 
— a  phase  most  important  in  relation  to  hygiene ;  but  in 
spite  of  this  and  although  the  statistics  are  only  approxi- 
mate, as  is  the  case  in  most  parasitological  work,  these 
data  have  attained  to  sufficient  proportions  to  justify  at 
least  a  beginning  in  the  matter  of  collating  and  general- 

'  There  are  certain  exceptions  to  this,  as  with  Nicoll's  (Proc.  Zool. 
«Sfoc.  London,  1912,  p.  858)  careful  search  for  trematodes  with  sieves,  but 
this  means  a  separate  research,  and  is  incompatible  with  the  all-round, 
general  policies  of  present  routine  laboratory  organization. 


616   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

ization.  At  any  rate  the  time  has  arrived  to  establish 
at  least  a  nucleus  for  the  accretion  of  data,  which  can  be 
later  subjected  to  conjSrmation  or  correction.  We  draw 
just  a  grain  of  comfort  from  the  knowledge  that  the  more 
fully  worked  field  of  human  parasitology  is  also  -vulner- 
able to  criticism  of  very  much  the  same  order. 

The  Value  of  Parasitological  Studies  in  Zoological 
Gardens. 

The  foregoing  chapters  have  made  clear  two  fields 
of  practical  usefulness  of  any  study  in  such  gardens. 
These — hygiene  in  relation  to  the  animals  and  comparison 
in  relation  to  human  beings — need  therefore  only  to  be 
mentioned  at  present  since  it  is  ob\T,ous  that  both  benefit 
by  our  parasitological  work.  But  there  is  yet  a  third — 
a  scientific  phase  of  parasitology  which  may  be  considered 
purely  academic.  It  consists  in  morphological  and  other 
studies  necessary  for  the  identification  of  the  parasite, 
the  determination  of  its  life  liistory,  etc.  These  last 
studies  may  still  in  a  restricted  sense  include  a  modicum 
of  the  practical  in  so  far  as  they  have  a  bearing  on  the 
disease  with  which  they  are  associated.  But  on  the  whole 
they  are  a  source  of  danger  for  us  since  such  things 
as  studies  on  the  finer  structures  of  worms,  taxonomic 
arrangements,  descriptions  of  new  species  of  commensals, 
etc.,  being  alluring,  are  likely  to  lead  one  so  far  afield  that 
eventually  an  attitude  of  stubborn  resistance  will  have  to 
be  assumed  in  order  to  conserve  that  precious,  volatile 
laboratory  asset — time — for  the  more  crying,  practical 
problems  ever  reaching  out  to  us. 

However,  in  parasitological  investigations  as  in  other 
scientific  work,  immediate  abstract  information  may  at 
some  time  prove  to  be  of  greatest  practical  value.  Thus 
for  example  if  we  can  discover  the  exact  facts  concerning 
one  phase  of  the  life  history  of  a  certain  parasite,  it  may 
be  possible  by  hygienic  measures,  to  break  the  cycle  of 
development  of  the  parasite  at  one  point  thereby  prevent- 


THE  ANIMAL  PARASITES  617 

ing  its  completion.  This  information  is  perhaps  obtained 
most  readily  in  experimentation  upon  the  role  of  lower 
animal  forms  in  the  pathogenesis  of  disease  but  where 
reliable  evidence  is  lacking,  help  may  be  had  by  compari- 
son with  others  in  the  same  taxonomic  group.  Undoubt- 
edly systematic  classification  will  go  far  to  help  solve 
many  of  these  riddles. 

Pathogenicity  of  Animal  Paeasites  in  General. 

The  first  question  which  arises  in  this  connection  con- 
cerns the  actual  ability  of  animal  parasites  to  produce 
disease  in  mid  animals.  At  once  it  will  be  seen  that  this 
must  be  a  relative  matter,  for  no  one  on  one  side  would 
contend  that  every  symbiont  in  an  animal  is  harmful — 
parasites  sensu  stricto — nor  on  the  other  that  none  could 
possibly  be,  i.e.,  that  all  are  always  commensals.  It  is 
evident  that  the  issue  boils  down  to  questions  as  to  the 
extent  to  which  they  are  harmful.  Before  attempting 
the  answer  let  us  consider  the  means  by  which  the  para- 
sites may  conceivably  produce  disease. 

Modes  of  Disease  Production  (Pathogenesis). 

The  medical  reader  is  familiar  enough  with  the  patho- 
genic powers  of  some  animal  parasites,  but  may  be  suffi- 
ciently interested  to  glance  over  specific  wild  animal 
instances  illustrating  them  while  they  are  being  listed  for 
those  less  familiar  with  this  subject. 

1.  Mechanical  Obstruction.   ; 

I  refer  here  particularly  to  simple  blockage  of  normal 
body  passages  as  the  result  of  bulk  or  mass.  This  occurs 
more  commonly  in  the  intestines  than  elsewhere  on  ac- 
count of  the  greater  frequency,  greater  numbers  and 
larger  size,  in  general,  of  parasites  inhabiting  this  tract. 
Thus,  we  have  recorded  a  liothrix  {Liothrix  luteus){2) 
where  the  combination  of  a  small  host  and  consequently 

(2)   Phila.  Zool.  8oc.  Rep.,  1920,  p.  28. 
40 


618   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

narrow  gut  and  comparatively  large  parasite  induced 
obstruction.  Plimmer(3)  records  microfilaria  clogging 
the  brain  capillaries.  Shipley  (4)  mentions  two  specimens 
of  Ascaris  lumbricoides  obstructing  the  nares  of  a  chim- 
panzee {Pan  niger).  Blockage  may  also  be  produced  sec- 
ondarily to  the  presence  of  the  parasite,  even  in  the 
absence  of  notable  numbers  of  them,  and  quite  apart  from 
the  element  of  verminous  bulk.  This  occurs  through  in- 
flammatory swellings  which  the  worms  excite.  We  saw 
many  serious  grades  of  this  in  our  spiroptera  epizootic, 
the  lumen  of  the  proventricle  being  narrowed  by  swelling 
of  the  mucosa  and  more  or  less  occluded  by  exudate  and 
necrotic  mucous  membrane. 

Yet  another  direction  wherein  a  mechanical  rationale 
pure  and  simple  obtains  is  by  the  production  of  divertic- 
ula. Worms  encysted  in  the  gut  wall  may,  by  weight 
alone  or  by  excitation  of  peristalsis,  cause  the  wall  to 
bulge  outwards  (or  inwards  even)  like  a  pocket.  Such  a 
diverticulum  has  been  noted  in  the  gut  of  a  Pale  Cebus 
{Cehus  ftavescens)  (5)  parasitized  by  acanthocephalus, 
but  in  this  case  there  were  adhesions  to  the  nearby 
stomach,  and  it  is  possible  that  in  this  individual  case  the 
diverticulum  was  a  traction  one,  i.e.,  pulled  out  by  the 
anchorage  of  adhesions  externally. 

2.  Mechanical,  Irritation. — In  those  instances  where 
inflammation  is  the  manifestation  which  reflects  the 
simple  mechanical  effects  of  parasites  it  will  be  difficult 
indeed  to  prove,  in  the  present  state  of  our  knowledge, 
that  it  is  not  rather  the  effect  of  associated  toxic  sub- 
stances or  excreta  elaborated  by  the  parasite.  But 
instances  of  a  purely  mechanical  irritation  there  must 
be,  although  one  can  scarcely  put  the  finger  upon  them 
and  say  that  this  or  that  individual  inflamed  mucosa  did 
not  become  so  from  a  toxic  cause.    Omitting  these  then, 

(3)  Proc.  Zool.  8oc.  London,  1910,  p.  134. 

(4)  Proc.  Zool.  Soc.  London,  1905,  p.  252. 

(5)  Phila.  Zool.  Soc.  Rep.,  1920,  p.  29. 


Fig.  71.— ACANTHOCEPHALUS  (THRKK  SPECIMENS)  PROJECTING  FROM  THE  IN- 
CISED INTESTINES  OF  A  PIGMY  MARMOSET.  COMPARE  THE  SIZE  OF  THE  PARASITES. 
WHICH  MAY  BE  DISTINGUISHED  BY  THEIR  ANNULATIONS.  WITH  THAT  OF  THE 
INTESTINES. 


Fig.  72. — BLOOD-RED  NEMATODES  PROTRUDING  FROM  FRONTAL 
SINUSES  OF  COMMON  OPOSSUM  (DIDELPHY8  VIRGINI  ANA).  IHE  SKULL- 
CAP HAS  BEEN  LIFTED  OFF  AND  THE  POSTERIOR  WALLS  OF  THE 
SINUSES  BROKEN. 


THE  ANIMAL  PARASITES  619 

the  more  certain,  purer,  more  unequivocal  examples  will 
be  those  where  physiological  processes  become  exalted  as 
the  result  of  the  parasitic  irritation.  An  example  in 
point  is  a  case  of  volvulus  in  a  Screech  Owl  {Otus  asio 
asio).{6)  Here  it  is  probable  that  the  parasites  excited 
the  gut  to  undue  peristaltic  action,  and  that  during  this 
process  it  became  twisted.  Worms  in  such  passages  as 
the  nose  and  nasal  sinuses  (I  have  seen  blood- red  filarias 
in  the  frontal  sinuses  of  an  opossum)  undoubtedly  pro- 
duce nervous  effects  by  their  presence  and  movements. 
Those  in  the  subcutaneous  tissue  (tilariae  of  wild  cats) 
probably  also  do  so.  It  is  difficult  to  judge  those  cases 
where  doubtfully  sensitive  parts  are  the  ones  affected. 
Probably  the  intestinal  and  intraperitoneal  worms,  and 
less  certainly  the  generally-migrating  ones  analogous  to 
Filaria  loa,  produce  no  nervous  effects  mechanically. 

3.  Production  of  Hemorrhages. — Hemorrhages  large 
enough  to  kill  suddenly  are  theoretically  possible,  since 
worms  occasionally  produce  aneurysms  which  may  rup- 
ture; we  have  seen  such  an  accident  in  a  Paradoxure 
{Paradoxurus  leucomystax).  But  certainly  it  is  the  long- 
continued,  wasteful  small  hemorrhages  that  are  impor- 
tant, inducing  an  anemia  often  of  severe  and  fatal  grade. 
The  hookworms  are  the  shining  offenders  here,  yet  we 
have  seen  very  much  the  same  effect  from  Acanthostoma 
in  the  intestine  of  monkeys.  (E'sophagostomum  has  also 
been  incriminated  at  the  London  Garden  in  young  Rhesus 
Macaques  {Macacus  rhesus)  {7)  where  the  young  forms 
of  the  parasite  did  the  damage  as  they  burrowed  into  the 
wall  of  the  gut. 

4.  Opening  up  Avenues  of  Infection. — This  may  be 
accomplished  either  by  passage  of  parasites  from  one 
position  normally  containing  bacteria  to  another  wliich 
is  susceptible  to  infection,  or  by  devitalizing  a  tissue 
which  is  ordinarily  resistant  to  infection;  i.e.,  creating  a 

(6)  Phila.  Zool.  Soc.  Rep.,  1921,  p.  31. 

(7)  Proc.  Zool.  Soc.  London,  1919,  p.  15. 


620   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

locus  resistenticB  minoris.  The  intestinal  tract  is  the 
most  common  organ  concerned,  but  the  illustrations  to 
follow  will  give  variety.  Thus,  the  mature  examples  of 
oesophagostoma  in  young  rhesuses  just  referred  to  above 
burrowed  into  the  gut  wall  and  led  to  both  local  and  gen- 
eral peritonitis.  In  one  of  our  **spiroptera"  parrots  the 
worm  had  passed  through  the  proventricular  wall  and  a 
chronic  fibrosis  resulted  around  it.  At  the  autopsy  on  a 
Rhesus  Macaque  Doctor  Fox  found  a  localized  abscess 
adjacent  to  the  gut  wall,  and  in  it  a  whipworm  was 
imbedded.  Passing  from  these  examples  of  intestinal 
worms,  I  can  mention  the  loss  of  a  valuable  Philippine 
Spotted  Deer  {Cervus  alfredi)  as  the  result  of  secondary 
infection  of  a  cysticercus  cyst  of  the  lesser  omentum 
which  led  to  a  nearby  peritonitis.  Lung  infections 
are  not  uncommon.  Murray (8)  records  that  forty-four 
out  of  eighty-five  young  rhesus  monkeys  dying  from  pneu- 
monia showed  an  acarian,  and  he  ascribed  the  pulmonary 
irritation  to  certain  crystals  in  the  excreta  of  the  mite.  I 
have  studied  a  case  of  bronchopneumonia  in  a  prairie  dog 
where  great  numbers  of  an  arachnid  were  present.  ,  The 
reports  of  the  London  Zoological  Society  are  replete  mth 
notes  of  round  worm  pneumonias  of  reptiles.  These  pul- 
monary cases  must  result  from  decreasing  of  tissue 
resistance  by  the  presence  of  the  worms,  and  are  easy  to 
understand,  much  more  so  than  the  intestinal  infections 
when  one  recalls  how  sensitive  lung  tissue  is  to  foreign 
bodies,  and  that  there  seems  to  be  no  indication  that  this 
tissue  becomes  accustomed  to  infestation  such  as  may  be 
argued  for  the  gut.  j  All  these  citations  must  convince  us 
that  parasites  are  most  important  predisposing  agents 
to  infection,  and  that  tliis  is  one  of  the  most  sinister 
phases  of  animal  parasitism.  . 

5.  Destruction  of  Tissue. — This  heading  does  not 
refer  to  the  comparatively  trivial  effects  that  accompany 
the  more  acute  inflammations   secondary  to   parasites, 

(8)   Proc.  Zool.  Soc.  London,  1919,  p.  15. 


THE  ANIMAL  PARASITES  621 

albeit  certainly  the  absorption  of  their  disintegrative 
tissue  products  has  some  effect  on  the  economy ;  but  our 
ideas  of  such  are  so  vague  as  to  justify  their  being  disre- 
garded here.  (What  I  refer  to  is  the  more  massive 
destruction  such  as  may  occur  in  the  blood,  for  instance, 
from  the  action  of  protozoa.  There  is  also  loss  of  mucosa 
in  those  chronic  inf estments  of  the  stomach  where  we  find 
excessive  fibrous  tissue  overgrowth.  The  most  striking 
example  of  tissue  destruction  we  have  seen  was  in  the 
cirrhotic  livers  of  prairie  dogs  affected  by  Eepaticola 
hepatica,  where  in  extreme  cases,  the  amount  of  function- 
ating liver  substance  was  reduced  to  a  very  smaU  fraction 
of  its  normal  bulk.  (9) 

6.  Toxins. — We  have  no  direct  evidence  to  offer  that 
noxious  products  of  parasites  are  concerned  in  producing 
disease  in  wild  animals.  The  local  effects  of  such  toxins 
are  not  distinctive  enough — individual  enough  to  toxins 
or  to  the  animal  body — to  separate  them  from  the  effects 
of  such  accompanying  factors  as  bacterial  inflamma- 
tions; nor  can  we  separate  the  general  effects  of 
these  toxins  from  what  might  have  been,  for  instance, 
the  effects  of  an  accompanying  anemia  of  hemorrhagic  or 
other  origin.  From  a  knowledge  of  what  happens  in 
human  prototypes  though,  there  is  scant  doubt  that  some 
one  of  the  multitudinous  species  must  be  capable  of  pro- 
ducing toxins,  but  just  which  varieties  are  concerned 
cannot  be  listed  by  anyone.  By  analogy  we  can  at  most 
only  suspect  the  hookworms  and  the  dibothriocephalidse. 
Under  this  same  category  of  the  toxins  come  the  worm- 
products  which  are  reputed  to  have  a  destructive  effect 
upon  the  digestive  enzymes  in  the  gastrointestinal  tract 
of  the  host,  and  which  would  thereby  interfere  with  the 
proper  assimilation  of  pabulum,  resulting  in  malnutrition. 
For  the  same  reasons  as  above  indicated  for  the 
toxins  one  is  unable  to  speak  for  or  against  these 
'  *  anti- enzymes. ' ' 

(9)   Phila.  Zool.  Soc.  Rep.,  1916-1921. 


622   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

7.  Preclusion  of  Nutrition.— This  must  be  a  very 
miimportant  phase  of  the  activity  of  intestinal  parasites, 
when  one  compares  the  bulk  of  food  which  passes  through 
the  bowel  and  the  average  number  of  worms  present ;  and 
the  same  holds  good  for  some  interstitial  parasites  like 
the  adult  filariaB.  Even  in  amazingly  hea\y  infestments 
of  the  intestines  one  will  be  constrained  to  dismiss  this 
idea  when  he  compares  the  bulk  of  parasites  with  that  of 
the  host,  and  recalls  what  the  physiologist  terms  the 
''factor  of  safety"  inherent  in  this  tract  as  elsewhere. 
But  in  the  case  of  blood  parasites  the  matter  may  be 
different.  Here  we  are  concerned  with  the  withdrawal  of 
refined  foodstuffs — those  which  have  been  worked  over 
and  over  by  subtle  internal  metabolic  processes ;  and  we 
are  not  so  sure,  especially  on  recalling  the  enormous 
numbers  of  parasites  usual  to  blood  infestments,  that 
there  is  the  capacity  on  the  part  of  these  internal 
processes  to  meet  increased  demands  that  we  count  upon 
for  the  intestinal  functions.  It  is  much  more  serious  to 
be  deprived  of  the  finished  product  than  of  the  crude 
because  it  means  the  undoing  of  "digestive"  work  all 
along  the  line,  from  gut  to  tissue  cell.  Furthermore,  a 
blood  infestment  guarantees  that  the  parasite  has  been 
feeding  upon  and  depriving  the  animal  of  the  precise 
foodstuffs  the  cells  require,  and  not  by  any  chance  upon, 
even  in  part,  intestinal  substances  that  were  wastes  or 
residues.  If  we  except  the  blood  parasites,  then,  it  seems 
safe  to  conclude  on  the  whole  that  the  amount  of  pabulum 
used  by  parasites  is  unimportant  to  the  animal. 

Having  reviewed  the  manner  in  which  parasites  may 
conceivably  be  harmful,  it  is  time  to  return  to  the  question 
of  the  actual  exercise  of  these  powers. 

The  older  appraisal  of  parasites  in  animals,  namely 
that  they  were  rather  innocent  of  disease  production,  was 
suggested  by  and  borrowed  from  the  veterinarian,  prob- 
ably being  engendered  in  him  by  their  frequency  in  what 
appeared  to  be  normal  domestic  specimens.  Yet  it  is  only 


THE  ANIMAL  PARASITES  623 

proper  to  add  that  one  of  our  former  pathologists,  and 
sometime  professor  of  veterinary  pathology,  Dr.  C.^  Y. 
White,  is  a  medical  man  and  is  of  much  the  same  opinion. 
Older  writers  regarded  worms  even  as  "guardian  angels" 
of  children.  Very  recently  Schwartz (10)  reviews  some 
work  in  this  connection  showing  that,  in  vitro,  some 
cestode  extracts  were  inhibitory  to  certain  bacteria  {B. 
anthracis,  B.  pyocyaneus  and  B.  dysenterice  Shiga). 
This  relationship  is  so  different  from  natural  conditions 
as  to  need  no  further  comment. 

At  the  London  Garden  the  view  appears  to  be  dif- 
ferent. In  the  1910  report  they  charge  five  deaths  against 
perforation  by  worms  of  the  stomach  and  intestines ;  in 
the  1911  report  they  record  giant  toads  dead  from  lung 
infestment;  in  1912  ''eighteen  cases  of  enteritis  were  due 
to  worms";  and  in  1917  they  mention  pneumonia  in  a 
toad  and  perforation  of  the  stomach  of  a  puma.  These 
reports  represented  evidently  the  more  striking,  unequiv- 
ocal examples  of  death  from  parasites  which  had 
outspoken  anatomical  expressions,  and  omitted  those  in 
which  the  more  subtle  agencies  of  parasitic  pathogenesis 
were  concerned.  Their  experience  has  apparently  been 
much  the  same  as  ours. 

The  ideal  approach  to  a  decision  in  reference  to  the 
importance  of  parasites  would  appear  to  be  a  mathe- 
matical one,  something  as  follows :  First,  to  determine 
what  species  infest  animals  and  how  commonly,  then  to 
decide  which  ones  are  pathogenic  and  tliirdly  to  estimate 
the  severity  of  the  disease  induced;  so  that  finally,  by  an 
analysis  and  comparison  of  the  three  results— a  com- 
parison and  analysis  judicial  in  the  broadest  sense— we 
might  hope  to  come  to  an  opinion.  Let  us  consider  the 
three  avenues  in  order.  At  the  first  glance  it  must  be 
evident  that  a  list  of  all  possible  parasitic  varieties  does 
not  exist  and  may  never  be  compiled.  The  most  that  can 
be  done  is  to  tabulate  the  findings  in  scattered  labora- 

(10)   Journal  of  Paras^it.,  June,  1921,  p.  194. 


624  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

tories,  data  usually  recorded  in  terms  of  the  individual 
observer's  studies  and  often  inadequate  to  give  the  com- 
piler all  the  facts  desired.  The  same  remarks  apply  to 
the  percentage  incidence  of  parasitism.  Not  to  prolong 
the  academic  discussion,  suffice  it  to  say  that  very  much 
the  same  obstacles  present  in  the  second  avenue — that  of 
pathogenicity  of  the  individual  species.  Our  own  data 
referring  to  this  second  heading  will  be  presented  later, 
but  after  the  failure  of  the  first  avenue,  the  second  and 
third  lose  greatly  in  value.  At  best,  statistics  can  be 
only  suggestive.  Unless  critically  and  suspiciously 
interpreted,  and  with  a  full  appreciation  of  their  limita- 
tions from  a  foreknowledge  of  the  way  in  which  they  were 
compiled,  they  would  only  delude  the  reader  and  offend 
science,  and  so  we  abandon  this  line  of  reasoning. 

At  present  the  best  results  of  the  study  of  patho- 
genesis by  animal  parasites  will  probably  be  reached  by 
a  combination  of  methods,  as  follows : 

1.  Direct.  How  commonly  do  we  see  clinical  symp- 
toms and  morbid  anatomical  changes  that  are  incontro- 
vertibly  due  to  the  parasite  ?  We  restrict  ourselves  here  to 
a  narrow  group  of  inf estments  indeed,  and  think  of  such 
diseases  as  trichosomiasis  in  prairie  dogs  and  spiroteria- 
sis  in  parrots. 

2.  By  comparison  with  analogous  infestments  of 
domestic  animals  and  man — more  thoroughly  studied  and 
therefore  more  accurately  appraised,  in  general,  as  to 
pathogenicity ;  a  comparison  from  the  standpoint  of  dis- 
ease production  rather  than  natural  habits  of  the  para- 
site. Example,  coccidiosis  and  hookworm  disease  in 
foxes  and  dogs. 

3.  By  inference  through  deduction.  This  is  the  most 
unsatisfactory  consideration  of  all,  and  should  be  well 
checked  up  and  discounted.  Here  we  would  evaluate  the 
known  propensities  of  the  parasite  first,  such  as  its  size, 
motility,  anatomic  position  in  the  host  and  the  general 
pathological  traits  of  the  genus  and  family  to  which  it 


THE  ANIMAL  PARASITES  625 

belongs,  etc.,  and  then  compare  these  verminous  properties 
with  those  of  the  host — its  size,  temperament,  physical 
stamina,  etc.  This  third  consideration  must  necessarily 
overlap  with  or  be  supplementary  to  the  first  two.  For 
example,  this  consideration  would  have  to  be  resorted  to 
in  many  cases  of  ascaris  infestment  where  anatomical 
changes  are  generally  not  demonstrable. 

Acting  on  these  three  considerations,  and  after  twelve 
years  of  observation  on  parasites  here  in  the  Garden,  a 
fresh  review  of  our  records,  and  a  recent  review  of  the 
accessible  relevant  parasitological  literature  I  have  come 
to  the  conclusion  that,  considering  wild  animal  collections 
the  world  over,  there  is  no  justification  for  an  unqualified, 
definite  answer  to  the  question  of  pathogenic  parasitism 
that  will  meet  all  conditions.  We  lack  data  on  too  many 
species  that  are  not  sufficiently  represented  in  collections 
or  indeed  not  represented  at  all.  It  is  the  liability  to 
infestment  of  each  order  or  family  of  beasts  that  will  have 
to  be  determined,  and,  depending  on  the  assortment  each 
garden  has  on  exhibition,  will  the  importance  of  parasites 
to  the  garden  as  a  whole  vary. 

Speaking  for  the  Philadelphia  Garden,  I  have  come  to 
the  conclusion  that  on  the  whole  parasitism  does  play  an 
important  part  of  our  animal  losses.  The  financial  loss 
which  could  be  charged  against  spiroptera  alone  is  in  the 
four  figures,  to  say  nothing  of  the  difficulty  of  replace- 
ment of  rare  species.  And  while  touching  the  financial 
phase  let  it  be  added  that  scientific  work  done  now,  it  must 
be  remembered,  is  not  restricted  to  the  present  time  or 
place,  but  is  to  be  measured  in  dollars  and  cents  with  the 
yard  stick  applied  to  the  future,  and  in  other  places  than 
that  where  the  initial  work  is  done.  Even  if  we  cannot 
answer  the  question  of  the  matter  of  importance  the  world 
over  we  can  guarantee  that  it  is  sufficiently  so  in  the 
Philadelphia  and  London  Gardens  to  warrant  a  rigid 
supervision  for  parasitism;  and  since  the  other  larger 
collections   are  probably  made  up  of  similar  animals, 


626   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

albeit  in  different  proportions,  we  surmise  at  least  that 
it  is  likewise  so  with  them. 

Importance  of  Parasites  in  Other  Fields. — In  addi- 
tion to  their  importance  to  exliibitions,  animal  parasites 
of  wild  animals  are  important  first  to  man.  The  animal 
hosts  may  serve  as  porters  of  infestation,  and  interfere 
with  attempts  at  eradication  of  the  disease.  The  experi- 
ence of  the  European  with  African  sleeping  sickness 
attests  to  this.  Not  to  go  farther  than  immediate  examples 
I  wish  to  note  in  this  connection  the  occurrence  in  this 
Garden  of  scabies  in  an  orang  which  was  transmitted  to  a 
keeper,  and  of  amebic  dysentery  in  monkeys.  Leiper(ll) 
has  called  attention  to  a  guinea-worm  in  a  leopard. 

Parasites  are  important  to  certain  wild  animal  indus- 
tries. The  ones  that  have  come  to  my  attention  are  the 
fur  seal  {Otoes  alaskamis)  industry  of  the  Pribiloff 
Islands  and  fox-farming  in  Newfoundland.  In  both  of 
these  instances  the  hookworm  was  concerned  and  entailed 
losses  of  thousands  of  dollars.  Lucas,  who  conducted  a 
United  States  Government  commission  to  the  seal 
grounds  and  after  whom  Stiles  named  the  parasite,  has 
left  very  full  notes  of  the  former  disease.  I  have  identi- 
fied the  same  infestment  in  a  young  California  hair  seal 
{Zalophus  calif ornianus)  which  was  born  and  died 
in  this  Garden.  This  indicates  that  the  parasite  might 
perhaps  be  found  farther  dowTi  the  Pacific  coast  than 
hitherto  suspected. 

To  hunters  parasitism  of  animals  must  be  important, 
but  to  an  unkno^vn  and  undoubtedly  unimagined  extent. 
The  grouse  plague  of  Scotland  (12)  is  an  example  to  point. 
Who  knows  but  that  the  disappearance  of  some  of  our 
game  animals,  particularly  birds,  was  not  due  more  to 
disease  than  to  the  ravages  of  man  I  There  is  at  least 
food  for  thought  here. 

(11)  Proc.  Zool.  Soc.  London,  1910,  p.  147. 

(12)  Fantham,  Proc.  Zool.  Soc.  London,  1910,  p.  672. 


THE  ANIMAL  PARASITES  627 

OCCUKEENCE    OF    AnIMAL    PaRASITES   IN    THE   WlLD. It 

would  be  unbelievable  that  parasitism  did  not  exist  in  the 
wild.  It  seems  proper,  however,  to  record  some  evidence. 
Diesing's  8y sterna  Hehninthum  is  replete  with  ref- 
erences to  Natterer's  Brazilian  expedition.  Nicoll  speaks 
of  a  German  expedition  to  Spitzbergen  in  1898,  and 
a  Swedish  one  to  Egypt  in  1901,  in  both  of  which  large 
numbers  of  parasitic  forms  were  collected.  Nicoll  (13) 
found  Trichosoma  hepaticum  in  a  hare  shot  in  the 
wild,  and  liver-flukes  (14)  in  a  kestrel  shot  on  the  coast 
of  Scotland.  Leiper(15)  found  nine  species  of  worms  in 
hippopotami  during  an  expedition  to  Uganda,  and  (16) 
states  that  thirty-seven  species  of  helminths  were  col- 
lected on  an  Antarctic  voyage  by  Surgeon  Atkinsons  In 
an  investigation  of  Grouse  disease  in  Scotland,  Fantham 
found  many  different  blood  and  intestinal  parasites.  \ 
Dr.  Charles  B.  Penrose  tells  me  that  all  of  the  white- 
tailed  deer  he  shot  in  the  valley  of  the  Swan  River,  Mon- 
tana, were  infested  with  liver-flukes,  so  much  so  that  the 
liver  was  literally  riddled  by  the  disease,  and  yet  the  deer 
were  fat.  The  black-tail  deer  of  the  same  valley  were  not 
thus  parasitized  and  were  not  as  fat.  In  our  own  Garden 
we  have  found  many  tapeworms  in  wild  cats  (17)  which 
had  been  too  recently  captured  for  the  worms  to  have 
developed  in  captivity.  Such  instances  might  be  still 
further  multiplied. 

A  more  important  consideration  is  the  fate  of  the 
parasites  thence  introduced  into  our  Garden.  Do  they 
disappear  of  themselves?  Naturally  we  can  never  make 
sweeping  predictions,  for  future  events  will  depend  upon 
the  life  history  of  the  individual  parasite  concerned.  But 
by  and  large,  once  introduced  it  is  better  to  assume  the 
attitude  of  pessimism,  and  resign  oneself  against  spon- 

(13)  hoc.  cit.,   1911,  p.  674. 

(14)  hoc.  cit.,   1915,  p.  87. 

(15)  Loc.  cit.,   1910,  p.  233. 

(16)  Loc.  cit.,   1914,  p.  222. 

(17)  Phila.  Zool.  Soc.  Rep.,   1912,  p.  40. 


628   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

taneous  disappearance  and,  what  is  worse,  realize  that  the 
parasitism  is  likely  to  become  indigenous.  We  have  sev- 
eral pieces  of  evidence,  however,  that  the  infestment  may 
occasionally  quite  disappear.  Thus,  I  have  seen  Coccid- 
ium  higeminum  spontaneously  disappear  from  a  Swift 
Fox  {Canis  velox)  and  Spiroptera  incerta  from  a  Macaw 
as  proven  at  autopsy.  Nicoll(18)  remarks  that  certain 
trematode  infestations  were  heavier  in  newly  arrived 
animals  than  in  ones  long  resident  in  the  Garden.  This  is 
.conceivable  on  the  basis  of  individual  worms  dying  out, 
i.e.,  fulfilling  their  life  spans  without  the  host  becoming 
reinfested  with  fresh  parasites.  Precise  information  on 
the  subject  is  supplied  by  Ackert(19)  who  found  that 
cestodes  disappeared  from  chickens  in  six  to  eight  months 
when  the  birds  were  confined,  i.e.,  protected  from  rein- 
festment.'i  Moreover,  it  is  known  that  worms  can  escape 
during  acute  infections,  the  infectious  state  of  the  economy 
producing  conditions  obnoxious  to  the  parasite.  We  hear 
of  many  instances  of  their  expulsion  in  human  feces  and 
vomitus  during  malaria  and  the  exanthemata  of  childhood 
and  know  of  similar  discharge  from  animals  during  the 
death  agony.  I  cite  these  data  largely  because  they  explain 
the  scarcity  or  absence  of  parasites  at  autopsy  in  animals 
which  were  known  to  have  been  clinically  infested. 

FREQUENCY  OF  PARASITISM  IN  WILD  ANIMALS 

There  can  be  little  doubt  that  mid  animals  are  more 
frequently  infested  than  man,  and  furthermore  with  a 
larger  number  of  parasites,  I  have  no  statistical  basis 
for  these  opinions — they  rest  on  personal  observations 
of  human  and  animal  autopsies,  and  reports  of  findings 
in  the  tropics  and  elsewhere.  They  have  therefore  but  the 
value  of  an  indi\ddual  opinion.  I  should  estimate  rather 
cautiously  that  ^dld  animals  are  infested  at  least  two  or 
three  times  as  frequently  as  man  and  much  more  heavily. 

(18)  Proc.  Zool.  Soc.  London,   1914,  p.  140. 

(19)  Jour.  Parasit.,  June,  1921,  Vol.  Vll,  p.  198. 


THE  ANIMAL  PARASITES  629 

The  first  step  in  the  discussion  of  the  incidence  of 
parasites  must  be  that  respecting  the  (host)  classes  and 
smaller  taxonomic  divisions — of  course  as  they  have  been 
studied  in  this  Garden.  Certain  statistical  limitations 
were  experienced  and  can  be  summarized  as  follows : 

Data  are  not  available  on  a  sufficiently  large  number 
of  animals  to  justify  conclusions  as  far  down  as  genera 
and  species,  except  for  such  commonly  and  generously 
exhibited  forms  as  monkeys  and  parrots.  I  have  there- 
fore in  tabulating  and  reviewing  our  records,  distributed 
the  animals  only  as  far  as  families — not  into  genera  and 
species.  The  table  (24)  to  follow  will  be  found  not  to 
contain  every  family  because  to  do  so  would  needlessly 
enlarge  it.  Accordingly  I  have  followed  the  policy  of 
only  indicating  those  genera  and  species  showing  either 
frequent  or  important  infestment.  I  shall  refer  to  those 
groups  later  as  *'  susceptible  "  groups.  If  no  family 
is  recorded  in  the  table  it  means  that  we  have  had  no 
important  numbers  of  inf estments  in  it.  The  ' '  remarks ' ' 
column  shows  the  individual  parasite  that  has  been  par- 
ticularly frequent  or  otherwise  important.  If  there  are 
no  remarks  it  means  that  the  species  of  parasites  found 
have  been  scattering. 

Results  of  Review  and  Tabulations. 

We  now  pass  to  an  analysis  and  discussion  of  the  find- 
ings brought  out  in  the  previously  mentioned  review  of 
our  records  and  in  Table  24.  Viewed  broadly  we  find 
that  there  is  a  wide  variation  in  the  susceptibility  of  dif- 
ferent families  to  infestment.  Those  that  are  susceptible 
may  be  located  by  consulting  the  table,  and  each  will 
therefore  not  be  separately  culled  out  and  subjected 
to  needless  repetition.  A  few  points  are  however  worthy 
of  separate  mention.  While  there  is  a  familial  or  generic 
susceptibility  within  certain  orders  it  is  unwise  to  gen- 
eralize too  broadly.    Thus  for  example  the  Corvidae  have 


630  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 


Table  24. 
Incidence  of  Parasites  in  Animal  Groups. 


Primates 

Cercopithecidae 

Sooty  Mangabey 

Cercocebus      fuligi- 
nosus 

Rhesus  Macaque 

Macacus  rhesus 
CalUtrichidse 

Marmosets 

Cebidae 

Squirrel  Monkeys 

Other  Cebus  Monkeys 

Lemures 

Carnivora 

FeUdse 

American  Wild  Cat.  .  . 
Fehs  ruffus 

Spotted  Wild  Cat 

Felis  ruflfus  texensis 

Canada  Lynx 

Felis  canadensis 

Lions 

Fehs  leo 

Ocelot 

Fehs  pardahs 
Canidse 

Gray  Fox 

Canis  cinereo  argen- 
teus 

Red  Fox 

Canis  vulpes  penn- 
sylvanicus 

Swift  Fox 

Canis  velox 

Gray  Wolf 

Canis  mexicanus 
Mustehdse 

American  Badger 

Taxidea  taxus 
Procyonidae 

Raccoon 

Procyon  lotor 
Ursidae 

Bears 


*538 
34 

60 

43 

8 

87 

86 

498 

28 

5 

10 

10 

15 

28 

17 

5 

18 

17 

42 
37 


9.4 
11.8 

10. 

16.3 

37.5 

11.5 

7. 

16.9 

40. 

80. 

40. 

30. 

33. 


12. 


40. 
11. 

41. 

5. 
16. 


Eight  had  Filaria  gracihs. 


Stomach  and  intestines, 
22;  Bronchi,  4;  Muscles, 
7. 

Ascarids  only. 

Ascarids  in  stomach  and 

intestines. 
Uncinaria. 


Cestodes. 

Uncinaria. 

Uncinaria. 
Ascarids. 


Physaloptera. 


Ascarids. 


♦This  figure  and  a  number  of  others  in  the  tables  do  not  correspond  with  those  in  other 
sections  of  this  book  because  certain  injured,  decomposed  and  newly  arrived  animals  were 
available  and  accepted  for  my  purposes,  but  were  objectionable  for  the  general  medical 
statistics  and  therefore  excluded. 


THE  ANIMAL  PARASITES 

Table  24  (Continued). 


631 


I 


Otariidse 

Hair  Seal 

Zalophus    californi- 
anus 

Rodentia 

Sciuridae 

Castoridae 

American  Beaver 

Castor  canadensis 

Hystricidse 

Canada  Porcupine.  .  . 
Erythizon  dorsatus 
dorsatus 


Hyraces 

Cape  Hyrax 

Procavia  capensis 

Ungulata 

Equidae 

Zebras 

Cervidae 

Axis  Deer. 

Cervus  axis 
Barasingha  Deer 

Cervus  duvanceli 
Eld's  Deer 

Cervus  eldi 
Fallow  Deer 

Cervus  dama 
Hog  Deer 

Cervus  porcinus 
Japanese  Sika  Deer.  . 

Cervus  sika  typicus 
Red  Deer 

Cervus  elaphus 
Elk 

Cervus  canadensis 
White  tailed  Deer .  .  . 

Mazama  virginiana 
Mule  Deer 

Mazama  hemionus 
Camelidse 
Llama 

Lama  glama 

Camels 

Suidae 


20 


198 
44 


17 


47 


365 


20 


14 


32 


17 


44 


5. 

16. 
9. 

23. 
36. 

28. 

12. 

100. 

17. 


62. 


14. 


Uncinaria. 


Scattered    through    four 
different  genera. 

In    three    cases    oxyuris 
and  flukes  in  cecum. 


Cestodes  8,  filaria  11,  oxy- 
uris 9,  in  peritoneal  cav- 
ity, also  intestine. 


Cestodes  in  bile  ducts. 

Nematodes,  intestine. 
C.  tenuicollis. 


Echinococcus  cysts. 


Trichocephalus. 
Echinococcus  in  lung  (2). 
Four  Cyst,  tenuicollis. 

Hydatid  cysts. 


632   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Table  24  (Continued). 


Animal 


Edentata 

Armadillos 

Marsupialia 

Didelphyidse 
Common  Opossum. .  . 
Didelphys     virgini 


Macropodidae 
Kangaroos  and  walla- 
bies  


'I 


16 

10 

175 

84 


70 


12.5 
20. 


48. 


Physaloptera,  38;  oxyuris, 
5;  cestodes,  5;  nema- 
todes in  lungs,  3;  cysts 
in  peritoneal  areolar  tis- 
sue, 2;  trematodes  in 
ileum,  1. 


Corvidse 

Common  Crow 

Corvus  brachyrhyn- 
chos  brachjThyn- 
chos 

Magpies 

Jays 


Pies,  choughs,  etc... 


Sturnidae 
Starlings . 


Turdidse. 


16 


35 


AVES 


Canaries. 


24 


12 


19 


44. 


64. 
55. 


33. 


Tropidocerca  and  occa- 
sional intestinal  cesto- 
des. Syngamus  in  crows. 
Few  filaria. 

Periproventricular  filaria, 
strongylus. 

There  is  a  striking  consis- 
tency of  infestment  in 
the  different  members  of 
Corvidse  both  as  regards 
degree  of  infestment  and 
species  of  parasite  pres- 
ent. 

Periproventricular  filaria 

largely. 
Periproventricular  filaria 
largely. 

Thrushes    and   Robins. 
None     in     American 
thrushes,    one    in    a 
robin. 
Finches.     Not  examined 
closely  at  autopsy,  but 
there  is  a  scattering  of 
periproventricular    fila- 
ria  and  intestinal    ces- 
todes through  most  of 
the  species. 

Were  free  from  parasites. 


THE  ANIMAL  PARASITES 

Table  24  (Continued). 


633 


AVES 

11 

II 

3  a 

1 
1 

4 

2 

50. 

30 

9 

30. 

142 

2 

7. 

*774 

124 

16. 

24 

5 

20. 

34 

2 

6. 

45 

4 

9. 

453 

65 

14.3 

121 

2 

1.6 

21 

6 

29. 

48 

21 

44. 

86 

12 

14. 

74 

13 

18. 

321 

69 

21.5 

26 

9 

34. 

62 

16 

26. 

164 

27 

16.5 

69 

17 

10. 

*201 

13 

6.7 

55 

4 

7.3 

44 

1 

2.3 

29 

1 

3.4 

73 

7 

9.6 

Picarise 
Picidse 

Woodpeckers 

Rhamphastidae 

Toucans 

Striges 

Psittaci 

Loriidse 

Lorys 

Cacatuidse 

Cockatoos 

Crested  Ground  Parra- 

keet 

Calopsitta  novse-hol- 
landise 
PsittacidsB 

Old  World  (Totals) . . . 
Undulated       Grass 

Parrakeet 

Melopsittacus     un- 

dulatus 

Pennant's  Parrakeet .  . 

Platycercus  elegans 

Rosehill  Parrakeet. .  .  . 

Platycercus  eximius 

Other  old  world  parra- 

keets 

Old     world     parrots, 

lovebirds,  eclectus  . . 

New  World  (Totals) . . .  . 

Macaws 

Conures 

Amazons 

Other  new  world  par- 
rots  

Accipitres 

Falconidse 

Buzzards 

Eagles 

Serpentaridse 

Vultures 

Miscellaneous 

*  For  foot  note  see  page  630 

41 


Spiroptera  largely. 
Remarkably  free  of  para- 
sites. 


3  spiroptera,  1  hemopro- 
teus,  1  intestinal  worm. 


2  spiroptera. 
4  spiroptera. 


1  spiroptera,  1  coccidium. 

6  spiroptera. 

20  spiroptera,  1  cestode. 

12  spiroptera. 

13  spiroptera. 

9  spiroptera. 

15  spiroptera,  1  hemopro- 
teus,  1  blood  larva. 

24  spiroptera,  3  nema- 
todes. 


7  spiroptera. 


4  were  blood  protozoa. 


634   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Table  24  (Continued). 


AVES 

Animal 

•ss 

II 

ll 

9  a 

1 

§ 

1 

Remarke 

Galli.. 

299 

95 
14 
70 

39 

♦163 

*38 

*41 

*21 

♦22 

♦105 

♦319 

48 

83 
188 
36 

42 

20 
3 
10 

7 

14 

7 
10 

3 

1 
21 
28 

7 

13 

8 

1 

14. 

21. 
21. 
14. 

18. 

9. 

18. 
25. 
14. 

5. 
20. 

8.8 
14. 

15.6 
4. 
2.8 

Phasianidae 

Quail 

Megapodidae 
Wild  Turkeys 

Columbse 

Fulicarise 

Heterakis. 

Intestinal  cestodes.    Coo- 

cidia  twice. 
Mostly  intestinal  cestodes, 

but  several  spiroptera. 

Alectorides 

Gavise. 

Steganopodes 

Herodiones 

Geese 

Parasites        scattering. 
Few  intestinal  cestodes. 

Ducks 

Struthiones. . 

*  For  foot  note  see  page  630. 

a  high  percentage  in  incidence  for  tropidocerca,  'syn- 
gamus  and  periproventricnlar  worms,  many  families  of 
Ungulata  harbor  echinococcus,  and  Carnivora  are  prone 
to  show  ascarids.    On  the  other  hand,  among  the  copious 
exceptions  to  this  may  be  cited  the  irregular  liability  to 
inf  estment  exhibited  by  the  Galli.  Four  varieties  of  these 
birds  are  represented  but  there  are  missing  such  impor- 
tant kinds  as  curassows,  guans,  guinea  fowl  and  peafowl.  I 
(Nor  do  all  members  of  a  genus  necessarily  show  the^ 
same  susceptibility,  and  the  heterakis  infestment  in  the 
pheasants  illustrates  this  matter  very  well.    It  was  limited 
almost  entirely  to  two  species — Amherst's  and  Golden, 
whereas  several  frequently  exhibited  species  showed  none.  ) 
The  following  table  brings  this  out  in  more  detail : 


THE  ANIMAL  PARASITES 


635 


Table  25. 
Heterakis  in  Pheasants. 


Species 


Golden  Pheasant  (Chrysolaphus  amherstise) . 
Amherst's  Pheasant  (Chrysolaphus  pictus). . 
Silver  Pheasant  (Gennaeus  nycthemerus) .  . . . 

Reeves'  Pheasant  (Phasianus  reevesi) 

Ringnecked  Pheasant  (Phasianus  torquatus) . 
Swinhoe  's  Pheasant  (Gennaeus  swinhoii) 


Enzootics  and  environment  played  no  part  in  the 
above  figures.  We  have  had  no  real  heterakis  enzootics, 
for  in  but  two  instances  did  three  heterakis  deaths  occur 
in  a  year,  and  two  deaths  per  year  have  occurred  in  but 
four  instances  in  the  past  twenty  years.  During  this  time 
there  have  been  sufficient  animals  on  exhibition  and  sub- 
jected to  autopsy  to  indicate  definitely  that  the  two  species 
named — Amherst's  and  Golden,  must  be  considered  as 
more  susceptible  than  the  other  varieties.  Nearly  all  of 
the  heterakis  in  quail  likewise  occurred  in  one  species — 
seven  of  the  ten  cases  occurred  in  a  total  of  twenty-three 
Scaled  Quail— but  in  these  birds  the  infestment  appeared 
in  enzootic  form  and  cannot  be  viewed  as  indicating  a 
preference  for  a  species. 

Psittaci  are  on  the  whole,  not  susceptible  to  worms. 
It  is  true  that  we  suffered  a  serious  outbreak  of  spiro- 
pteriasis  a  few  years  ago,  but  if  we  consider  this  a 
closed  chapter  we  can  accept  the  above  generality  as 
stated.  Among  774  parrots  autopsied  we  have  encoun- 
tered but  one  cestode  and  three  intestinal  round  worms. , 

The  deer,  likewise,  are  singularly  free  from  intestinal 
parasites.  I  gave  the  detailed  records  of  these  animals 
in  Table  24  to  emphasize  the  scarcity  of  parasites  even 
when  fairly  numerous  specimens  had  been  available  for 
examination. 

Other  interesting  features  in  the  table  are  the  out- 
standing infestments  of  squirrel  monkeys  and  marmosets 
among  the  monkeys,  of  gastric  and  intestinal  worms  in 
the  wild  cats,  and  intestinal  worms  in  the  zebras. 


636  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 


The  foregoing  has  had  to  do  with  parasitism  from 
the  standpoint  of  the  host.  The  next  phase,  that  of  the 
individual  parasite  itself,  interests  more  the  strict  parasi- 
tologist than  the  general  zoologist;  however,  both  will 
see  how  it  may  have  a  very  practical  value. 

Table  26. 

Distribution  of  Parasitic  Cases  According  to  Parasitic  Groups. 


Nematodes 

Spiroptera 

Filaridae 

Ascaris 

Physaloptera 

Uncinaria 

Tropidocerca 

Heterakis 

Trichocephalus 

Syngamus 

Trichina 

Hepaticola 

Other  Miscellaneous. 

Total  Nematodes. . .  . 
Cestodes 

Echinococcus 

Cysticercus 

Taenia 

Miscellaneous 

Total  Cestodes 

Trematodes 

Acanthocephalus 

Protozoa 

Arthropods 

Unclassified 


183* 

145 

138 

30 

28 

25 

23 

22 

11 

9 

2 

2 

4 

165* 
9 
7 
4 
3 


622 


188 
22 

4 
14 

6 
34 


Grand  Total. 


890 


*  Not  generically  diagnosed. 

Incidence  According  to  Parasitic  Groups. 

Inasmuch  as  it  has  been  physically  impossible  to  de- 
termine specifically  and  classify  efficiently  the  accumu- 
lations of  verminous  material  from  our  autopsies  I  will 
not  be  able  to  tabulate  parasitic  groups  even  as  closely  as 
I  did  in  the  * '  animal  host  ' '  table.  Nevertheless  sufficient 
has  been  done  to  illuminate  in  part  certain  phases  of 
parasitism  and  to  prevent  a  summary  dismissal  of  the 
subject.  Reviewing  our  cross-index  I  have  distributed 
the  data  into  the  following  Table  26,  the  parasites  being 


THE  ANIMAL  PARASITES  637 

listed  in  the  order  of  their  frequency.  It  may  serve  only 
as  a  panorama  of  the  situation,  inasmuch  as  determinative 
study  of  a  group  amounts  to  a  research  in  itself,  and  the 
multiplicity  of  them  precludes  a  consistent  study  of  every 
one.  The  data  are  based  upon  ''  cases  of  parasitism." 
That  is,  each  and  every  worm  species  occurrence  has  been 
counted,  regardless  of  whether  it  was  the  same  species 
that  has  been  concerned  over  and  over  again,  or  in  dif- 
ferent anatomical  positions  (of  different  individual  hosts, 
of  course)  or  whether  it  was  in  association  with  other 
parasites. 

Analysis  of  Table  26. 

There  is  a  grand  total  of  890  cases  of  animal  parasit- 
ism embraced  in  the  above  table,  which  is  a  sufficiently 
large  number  to  give  representative  value  to  some  phases 
of  the  analysis. 

I  In  the  first  place  nematode  worms  occur  about  three 
times  as  frequently  as  all  other  forms  of  parasites.  In 
gardens  where  spiroptera  has  not  figured  so  largely  the 
proportion  might  be  reduced  to  about  two  to  one.  Ces- 
todes  rank  a  poor  second,  trematodes  a  worse  third,  and 
acanthocephali  a  very  bad  last.  This  order  agrees  with 
our  figures  of  1913(20)  and  with  the  small  series  of 
Nicoll.(21)  The  latter  worker  found  that  the  order  was 
not  changed  when  pains  were  taken  to  include  also  such 
smaller  worms  as  could  only  be  obtained  from  the  host  by 
using  sieves,  etc.  Cestodes  were  not  likely  to  be  over- 
looked, but  very  small  trematodes  and  nematodes  were 
easily  passed  over. 

ViscEEAL  Distribution. 

As  to  the  individual  organs  which  are  most  commonly 
parasitized  our  records  show  that  with  Aves  as  well  as 
Mammalia  the  intestines  are  the  parts  most  commonly 
affected.    The  stomach  ranks  second  for  both-\-the  pro- 

(20)  Proc.  Acad.  Nat.  8ci.  Phila.,  March,  1913,  p.  127. 

(21)  Proc.  Zool.  Soc.  London,  1912,  p.  858. 


638  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

ventricle  rather  than  the  gizzard  of  birds  corresponding, 
parasitologically  speaking,  to  the  stomach  of  mammals. 
We  have  found  but  one  parasitic  species  in  the  gizzard 
of  birds,  i.e.,  immature  forms  of  Spiroptera  incerta  lying 
under  the  chitinous  lining  of  the  gizzard  and  only  dis- 
coverable after  the  lining  has  been  peeled  off.  The  peri- 
toneum comes  third  (air  sacs  of  birds)  due  to  the  presence 
of  filaridae,  and  the  blood  fourth  for  the  same  reason."  It 
is  to  be  emphasized  that,  in  our  data,  identical  organs 
of  mammals  and  birds  should  be  about  equally  liable  to 
infestment  with  the  possible  exception  of  the  lungs.  But 
in  view  of  the  small  number  of  cases  available  there  is  no 
justification  for  speculating  about  the  reason  for  this 
last  difference,  albeit  the  radical  difference  in  the  anatomy 
of  the  two  classes  is  very  inviting. 

Now  that  our  spiroptera  enzootic  has  subsided,  the 
order  above  given  will  be  changed,  and  in  view  of  like 
disturbing  factors  other  gardens  should  not  expect  the 
same  order  to  hold  invariably  for  their  collection,  since 
their  enzootics  will  depend  somewhat  on  the  preponder- 
ance of  animals  of  one  or  another  family  which  are  likely 
to  compose  their  exhibits.  A  single  such  enzootic  may 
suffice  to  disarrange  the  whole  fabric,  and  if  two  or  three 
are  taken  into  account  the  order  of  organ  involvement 
can  be  quite  disrupted.  To  attempt  to  construct  statisti- 
cally an  '*  order  of  frequency  involved  "  which  would 
stand  for  every  garden  would  only  lead  to  interminable 
adjustments  on  the  basis  of  animals  exhibited  and  of 
parasitic  enzootics,  so  that  I  have  finally  been  reduced  to 
a  combination  of  our  Garden  statistics  and  the  blood- 
parasitic  ones  of  the  London  Garden.  Doing  this  I  have 
arranged  in  Table  27  the  frequency  of  organ  involvement 
as  follows  and  estimated  the  percentage  of  animals  in- 
fested. These  figures  are  computed  upon  a  different  basis 
from  that  of  Table  24.  They  naturally  cover  all  animals 
and  not  the  *'  susceptible  "  ones  as  in  Table  24. 


Fig  73.— hugely  DISTENDED  PROVENTRICLE  OF  PARROT  DYING  WITH 
SPIROPTERIASIS.  COMPARE  ITS  SIZE  WITH  THAT  OF  THE  HEART  WHICH  IS 
ABOVE  AND  TO  THE  LEFT,  AND  THAT  OF  THE  GIZZARD  BELOW  AND  TO 
THE   LEFT. 


THE  ANIMAL  PARASITES 

Table  27. 

639 

Mammalia 

Aves 

per  cent. 

9.0 
3.7 
2.3 
1.5 
1.0 
1.0 
0.5 

Blood 

per  cent. 

6.5 

Intestines 

3.5 

Proventricle 

1.7 

Blood 

Air  sacs 

1.3 

0.3 

0.3 

0.4 



Total 

20.0 

Total 

14.0 

/ 


(The  effect  of  this  is  at  first  sight  startling  in  that  it 
places  the  blood  parasites  of  birds  so  far  in  the  fore,  but  it 
must  be  at  once  recalled  that  the  inquiries  upon  the  blood 
parasites  were  much  more  searching — microscopic,  than 
in  the  case  of  the  other  organs.  If  similar  methods  were 
applied  to  the  others  their  percentage  of  parasitism  might 
be  notably  raised — particularly  that  of  the  intestines.  ) 

Special  Parasitologic  Considerations. 

At  this  point  the  statistical  considerations  of  parasit- 
ism will  give  way  to  descriptions  of  certain  specific  inf  est- 
ments  that  have  given  us  more  or  less  concern. 

The  occurrence  of  single  parasitic  varieties  or  of  well 
known  species  in  an  isolated  host  may  occasionally  be  of 
practical  importance,  but  usually  they  amount  to  little 
more  than  an  academic  study,  whereas  the  repeated  dis- 
covery of  single  parasitic  kinds,  or  inf  estment  of  similar 
hosts,  especially  when  grouped,  raises  the  matter  to  a 
very  practical  level  demanding  attention.  Such  findings 
being  not  infrequent  in  our  experience,  it  has  been  possi- 
ble to  study  our  material  in  a  manner  designed  to  show 
the  frequency  of  various  parasites  in  a  certain  host,  the 
susceptibility  of  certain  animals  to  parasites  in  general 
and  the  inf  estment  of  dissimilar  hosts  by  the  same  para- 
site.   The  more  important  of  these  now  follow. 


640  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

Avian  Spibopteriasis. 

'  This  disease  concerned  parrots  particularly  but  tou- 
cans, pigeons,  and  such  widely  separated  species  of  birds 
as  the  starling,  quail,  thicknee  and  barbet  have  been  occa- 
sionally affected.  To  the  naked  eye  the  parasite 
resembles  the  human  hookworm,  but  differs  in  location, 
being  a  resident  of  the  proventricle  where  it  produces 
a  swelling  of  the  mucosa  which  interferes  with  the  passage 
of  food.  Up  to  a  hundred  worms  may  be  present  in  the 
one  bird,  and  immature  fonns  are  occasionally  found 
under  the  chitinous  lining  of  the  gizzard.  The  parasite 
burrows  into  the  mucous  membranes,  occasionally  pene- 
trates quite  through  the  wall  into  the  air  sacs,  and  on 
one  occasion  induced  an  adenomatous  hyperplasia  of  the 
mucous  membrane,  and  an  adjacent ' 'peritonitis. ' '  Mucus 
is  sometimes  present  in  the  droppings.  Death  may  occur 
either  acutely,  or  with  emaciation.  Spiroptera  incerta 
Smith (22)  is  the  common  parasitic  species  of  parrots,  but 
I  have  found  at  least  one  other  as  yet  unidentified  species 
in  the  toucan,  and  there  are  proba.bly  more.  In  the 
eight  year  period  1906-1913  from  25  to  50  per  cent. 
of  our  dead  parrots  showed  this  parasite  every  year, 
the  total  loss  being  113  birds  for  this  period — a  most 
important  inf estment.   ' 

We  approached  the  problem  by  diagnosing  and  isolat- 
ing the  infested  birds  through  a  microscopic  examination 
of  droppings,  finding  that  by  boiling  the  droppings  in  5 
per  cent.  NaOH  solution  we  clarified  them  and  made  ex- 
amination easier  and  more  certain  without  at  the  same 
time  destroying  the  parasitic  ova.  The  result  of  the  ex- 
amination of  all  our  parrots  was  the  isolation  of  14  per 
cent,  of  the  parrot  population ;  and  as  these  died  off  the 
diagnosis  of  inf  estment  was  found  confirmed  at  autopsy 
in  every  case.  The  parrot  house  was  thoroughly  reno- 
vated and  no  newly  arrived  parrots  were  admitted  until 
after  quarantining  and  examining  droppings  for  ova. 

(22)   Proc.  Acad.  Nat.  8ci.  Phila.,  1913,  p.  133.  ' 


4^      ^  '        fjv\i-* 


Fig.  74.— histologic  SECTION  THROUGH  PROVENTRICULAR  WALL  OF  PAR- 
ROT. SHOWING  SECTIONS  OF  SPIROPTERA  IN  THE  LUMEN  AND  MUCOSA.  THERE 
IS  SOME  GLANDULAR  HYPERPLASIA  (ADENOMATOID)  AND  NECROSIS  OF  THE 
LUMINAL   PORTIONS   OF  THE  MUCOSA. 


Fi<;.  75.— INFLAMMATORY  ROUND-CELL  INFILTRATION 
AROUND  NERVE  TRUNK  IN  WALL  OF  PROVENTRICLE.  PARROT 
DEAD  WITH   SPIROPTERIASIS. 


THE  ANIMAL  PARASITES  641 

The  toucans  and  other  species,  being  housed  elsewhere, 
were  not  quarantined.  Following  this,  we  were  gratified 
to  experience  no  more  spiroptera  deaths  in  parrots  for 
seven  years.  Then,  in  1920  and  1921,  a  new  outbreak 
occurred  in  four  toucans  and  several  other  scattering 
species,  including  two  parrots;  but  none  of  these  came 
from  the  main  parrot  house  and  probably  represented  a 
fresh  importation.  We  attempted  to  cure  the  isolated 
verminous  birds  by  medication  but  were  unsuccessful. 
Likewise  attempts  at  determining  the  life-cycle  of  the 
parasite  brought  us  no  farther  than  that  the  ova  devel- 
oped larvae  in  moist  sand  in  six  days.  Feeding  of  ova, 
freshly  passed  and  larvated  did  not  produce  infestment 
in  parrots  or  pigeons.  On  the  whole  we  can  quote  our 
experience  with  spiroptera  as  a  most  satisfactory  exam- 
ple of  the  value  of  hygiene  and  as  a  result  which  could 
never  have  been  accomplished  by  medication. 

Hepaticola  (Trichosoma)Hepatica  in  Prairie  Dogs. 

Bancroft  (23)  and  Hall  (24)  have  given  us  details  con- 
cerning this  parasite  and  the  disease  it  causes.  It  is 
threadlike,  several  inches  long,  and  permeates  the  livers  of 
the  gray  rat,  white  rat  and  wild  hare.  (25)  We  first  saw  it 
in  the  more  or  less  cirrhotic  livers  of  several  prairie  dogs ; 
later  we  observed  it  in  a  beaver  and  the  gray  rats  of 
the  Garden.  In  the  prairie  dogs  and  beaver  the  liver 
resembled  that  of  fatty  cirrhosis  and  was  so  considered 
on  naked  eye  examination  at  our  first  autopsy.  We 
were  only  set  right  when  we  came  to  the  histological 
examination.  It  was  remarkable  how  well  conditioned 
some  of  the  prairie  dogs  were  in  in  the  face  of  very  exten- 
sive liver  destruction;  but  on  the  other  hand  some  were 
emaciated  and  a  few  of  the  spontaneously  diseased  showed 
at  autopsy  an  enormous  ascites.  The  outstanding  features 
at  autopsy  were  the  large  size  of  the  liver  and  its  pallor 

(23)  Proc.  Roy.  Soc.  N.  So.  Wales,  Sydney,  Vol.  27,  pp.  86-90,  1893. 

(24)  Proc.   U.  8.  Nat.  Mus.,  Wash.,   D.  C,  Vol.   50,   1916,  p.   31. 

(25)  Proc.  Zool.  Soc.  London,  1911,  p.  674. 


642   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

and  hardness;  and  fine  yellow  lines  could  sometimes  be 
made  out  twisting  over  the  surface. 

The  disease  affects  wild  rats  differently  from  prairie 
dogs.  In  both  the  spontaneous  and  experimental  disease 
the  infestment  was  insignificant,  amounting  to  perhaps 
three  or  four  foci  the  size  of  a  split  pea  near  the  anterior 
margins  of  the  liver.  Diagnosis  may  be  easily  confirmed 
by  crushing  the  yellow  infested  portions  of  the  liver 
between  glass  slides  and  examining  microscopically 
for  ova. 

"We  have  seen  such  a  small  number  of  cases  of  this 
disease  because  so  few  prairie  dogs  reach  the  autopsy 
table,  yet  there  must  be  some  important  mortal  factor 
in  our  prairie  dog  enclosure,  for  the  Superintendent 
states  that  the  population  there  does  not  increase  in  spite 
of  the  frequent  births  and  additions  from  dealers.  The 
animals  almost  invariably  die  under  ground  and  their 
bodies  are  not  recovered. 

In  order  to  test  out  the  origin  of  the  infestment  we 
trapped  two  of  our  exhibition  specimens,  and  the  liver  of 
both  was  found  infested  on  surgical  examination  whereas 
six  newly  purchased  ones  had  normal  livers.  The  latter 
were  secured  fresh  from  their  native  habitat  in  the  West, 
and  their  livers  were  examined  through  long  surgical 
incisions  and  found  free  of  infestment.  Later  we  fed  the 
ova  (embryophores)  from  rat  livers  to  these  prairie  dogs 
and  on  destroying  them  found  them  infested.  We  were 
also  successful  in  transmitting  the  disease  in  the  oppo- 
site direction,  i.e.,  from  prairie  dog  liver  to  white 
rat.  From  all  this  we  feel  sure  that  the  prairie  dog 
disease  in  our  Garden  was  transmitted  from  the  rat  and 
that  here  is  another  reason  for  rat  extermination  in  a 
zoological  garden. 

The  adult  Hepaticola  hepatica  of  prairie  dogs  I  have 
not  seen  in  sufficient  entirety  to  compare  with  the  rat  spe- 
cies and  therefore  cannot  affirm  that  the  two  are  identical 
species.  It  is  presumably  like  that  of  the  rat,  being  thread- 


nr.    76— OVA    OI     HtPATICOLA    HIPAflCA    IN    lUFR    Ol     ''^ARIH  •  "<-       '  ^^^ 

HAM    BIPOI  \R  OPtMNCb       IHhRF    IS  DI  bl  RUC  I  ION   OI    '-' V^    ' '""^  vo  mPMH  1  ^M 

INHAMMAIORY      REACTION      OF     ChU.ULAR      CHARAClhR       HLT       NO  IMPORI^Nl 
FIBROSIS. 


r  i  I  ^1  I  ^1  I  "^1  I  ^1  I  ^1  1 1  I  ®l  1  ^ 

'iiiiiiiiiliiiiliiiiliiiiliiiiliiiiliiiiliiiilitiiliiiniiiiltiiiliiiiliiiiliiiil^ 


Vu..  77.— r\ti\  ar:  A  >\iriHi  cdiiii)  i\  i\  ik  \iii:i' \  i  k'  iiii.i:  dicis  ok 

GIRAFFE.      Nf)TE   MARKKI)    TKRIDI  CI  AL   FIBROSIS    IN    I  H  K    N  KI<;HBORHOOI)    OF 
THE  PARASITES. 


THE  ANIMAL  PARASITES  643 

like  and  most  difficult  to  separate  from  the  liver  substance 
through  which  it  ramifies.  At  maturity  it  dies  and  dis- 
integrates, leaving  the  ova  distributed  more  or  less 
in  tracts  through  the  liver  substance,  so  that  we  are 
limited  to  a  certain  period  wherein  to  obtain  the  mature 
form.  The  ova  are  not  passed  into  the  intestine,  but  re- 
main in  situ,  just  as  in  the  case  of  hydatid  disease,  and 
therefore  diagnosis  cannot  be  achieved  by  examination 
of  feces.  For  the  disease  to  be  transmitted  the  host  must 
die  and  its  carcass  be  eaten  or  otherwise  so  disintegrated 
that  the  ova  are  distributed  abroad.  Another  interest- 
ing observation  is  the  long  incubation  period  of  the 
ova.  Confirming  Bancroft,  we  found  that  the  ova  only 
became  larvated  after  they  had  lain  in  water  at  least 
three  months. 

Hookworms. 

These  important  parasites  have  been  taken  from 
several  foxes:  Gray  Fox  (Canis  cinero  argenteus),  Arc- 
tic Fox  {Canis  lagopus),  Swift  Fox  {Canis  velox),  Red 
Fox  {Canis  vulpes  p ennsylv aniens) ,  a  Gray  Wolf  {Canis 
mexicanus),  divers  members  of  the  Felid£e-Eyra  {Felis 
eyra),  Jaguarundi  {Felis  jaguarundi),  American  Wild 
Cat  {Felis  ruff  us),  Spotted  Wild  Cat  {Felis  ruff  us 
texensis),  Ocelot  {Felis  pardalis),  from  two  Giraffes 
{Giraffa  camelopardalis,  Giraffa  capensis),  a  Malayan 
Tapir  {Tapirus  indicus),  and  a  young  California  Hair 
Seal  {Zalophus  calif ornianus).  It  has  been  a  most  seri- 
ous infestment  in  American  wild  cats  {Felis  ruffus  and 
Felis  ruffus  texensis) — animals  which  generally  also  har- 
bor other  species  of  worms.  In  view  of  the  petechial 
hemorrhages  of  the  intestines  and  analogous  circumstan- 
ces in  dogs  and  human  beings,  it  must  be  conceded  that 
this  worm  is  pathogenic. 

At  this  point  it  is  fitting  to  note  the  infestment  as  it 
affects  hair  seals.  The  parasite  concerned,  Uncinaria 
lucasi,  has  long  been  a   scourge   among  the   fur  seals 


644   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

{Otoes  alaskanus)  of  the  Pribiloff  Islands.  Its  punctures 
are  bloodless,  being  signalized  instead  by  small  edem- 
atous plaques  in  the  intestinal  mucosa.  The  animal  we 
autopsied  was  a  young  California  Hair  Seal  born  in  the 
Garden,  and  is  singularly  the  only  hair  seal  in  which  we 
have  seen  it.  The  natural  habitat  of  the  hair  seal  is  the 
coast  of  California  which  means  that  the  range  of  U. 
lucasi  may  extend  farther  southward  than  at  first  sus- 
pected.    We  have  none  of  the  northern  variety. 

I  point  out  two  giraffe  cases  only  because  they  are 
unique  as  to  the  organ  (liver)  affected.  So  far  as  I  know, 
mature  hookworms  have  never  been  reported  from  other 
organs  than  the  intestines. 

From  the  prophylactic  standpoint  it  mil  be  advisable 
to  have  as  little  moist  earth  as  possible,  particularly 
sandy  ground,  in  and  around  the  enclosures  for  the  above 
mentioned  susceptible  animals  because  it  is  in  such  soil 
that  the  earlier  stages  of  the  life  cycle  of  the  parasite 
are  passed. 

We  have  never  found  any  of  the  human  hookworm 
species  in  our  animals,  but  it  must  be  recognized  that 
transmission  is  possible  to  a  certain  degree.  Anchylostoma 
ceylanicum  Lane  (26)  was  found  in  man,  cats,  dogs,  and 
a  lion;  Leiper(27)  reports  A.  duodenale  in  a  dog,  and 
von  Linstow(28)  states  that  the  latter  parasite  also 
occurs  in  the  chimpanzee. 

Amebic  Dysentery  in  Monkeys. — We  recently  lost 
six  monkeys  in  a  small  outbreak  of  this  disease — four 
black  spider  monkeys  {Ateles  ater),  a  Pinche  marmoset 
{Leontocehus  edipus),  and  a  woolly  monkey  (Lagothrix 
lagotricha).  Except  for  non-characteristic  looseness  of 
stools,  there  were  no  symptoms  until  the  usual  terminal 
lethargy  set  in.  Li\^ng  amebae  were  found  in  feces.  At 
autopsy  only  the  colon  was  found  to  be  anatomically 

(26)  Indian  Med.  Gaz.,  June,  1913,  p.  217. 

(27)  Jour.  Trop.  Med.  Etc.,  London,  1913,  XVI,  p.  334. 

(28)  Am.  Med.  Phila.,  V.  6   (16),  1903,  p.  611. 


Fig.   78.— microscopic  SECTION  OF  LIVER  OF  GIRAFFE.   SHOWING  SECTIONS  OF  UNCI- 
NARIA  SMITHI   IN    BILE-DUCT  AND   MARKED   FIBROSIS  AROUND  THE  DUCT. 


Fig.   79.— colon   OF  MONKKV   OVINC 


KI.KVATKI)   SLOUGHS 


THE  ANIMAL  PARASITES  645 

affected.  It  was  hugely  distended,  fully  an  inch  in 
diameter,  and  there  were  numerous  confluent  ulcers  of  the 
mucosa  covered  by  a  thick  slough.  The  liver  showed  no 
abscesses.  In  the  histological  sections  amebae  were  found 
in  the  interstices  of  vital  gut  tissue  just  as  they  are  in 
corresponding  human  lesions.  I  have  not  diagnosed  the 
species  yet,  but  can  vouch  that  it  is  not  Endameha  his- 
tolytica or  coli. 

According  to  Leidy  's  recommendation,  grated  nutmeg 
was  administered  and  was  followed  by  an  improvement 
in  symptoms.  The  animals  became  brighter  and  the  stools 
firmer,  but  the  amebae  were  not  eradicated.  Emetin 
hypodermically  and  by  mouth  had  no  obvious  effects  on 
the  disease  or  the  amebae.  One  monkey  thus  treated  with 
nutmeg  recovered,  but  died  the  next  year  of  another  affec- 
tion and  disclosed  the  scars  of  the  old  ulcers  in  the  colon. 
Our  experience  with  this  disease,  however,  is  not  unique. 
At  Washington,  D.  C.,(29)  eight  spider  monkeys  were 
affected,  and  sporadic  cases  come  to  light  from  the  West 
Coast  (30),  Manila,  Khartoum  and  Ceylon.  Prowazek's 
report  concerned  a  young  orang(31).  Liver  abscesses  in 
addition  to  the  intestinal  lesions  were  found  by  three 
different  observers. 

As  to  the  transmissibility  of  monkey  amebiasis  to 
man,  reporters  are  divided.  Both  sides  are  probably 
right,  in  as  much  as  Endameha  histolytica  was  concerned 
in  some  cases  and  non-human  species  in  others.  It  is 
an  infestment  to  be  feared,  and  calls  for  examination  of 
stools  from  such  newly  arrived  animals  as  are  known  to 
be  susceptible  (spider  and  woolly  monkeys,  orangs). 

Parasites  of  Marmosets  and  Squirrel,  Monkeys. — I 
give  a  special  place  to  this  subject  because  Table  24  shows 
that  these  monkeys  are  so  commonly  infested  and  because 
they  are  so  commonly  used  as  household  pets.    In  this 

(29)  Eichhorn  and  Gallagher,  Jour.  Inf.  Dis.,  XIX,  No.  3,  Sept., 
1916,  p.  395. 

(30)  Macfie,  Ann.  Trop.  Med.  and  Parasit.,  1915,  9,  p.  507. 

(31)  Arch.  f.  Protistenk,  Jena,  V.  26   (2),  22,  July,  p.  241. 


646   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

connection  the  questions  arising  are,  first,  whether  the 
infestment  is  a  menace  to  life,  and  second,  whether  it  is 
existent  outside  the  Garden  or  only  acquired  here.  The 
following  lists  set  forth  the  parasitic  status  as  sho\vn  at 
autopsy.  The  figures  indicate  how  long  the  animal  lived 
in  the  Garden : 


Marmosets 

Squirrel  Monkeys 

Infested 

Not  infested 

Infested 

Not  infested 

1  day 

1  day 

2  months 
6  months 

12  months 
12  months 

12  months 

13  months 

6-15  days         (  4  animals) 

1  month         (  6  animals) 
3-5  months    (  9  animals) 

6  months        (  2  animals) 

7  months       (  2  animals) 

8  months        (  1  animal) 

9  months        (  2  animals) 
10  months        (   1  animal) 
12  months        (  1  animal) 

14  months        (  1  animal) 

15  months        (  1  animal) 

17  months       (  1  animal) 

18  months        (  2  animals) 

20  months       (  1  animal) 

21  months        (  1  animal) 

2  days 

14  days 
26  months 

3  months 

3  months 
5  months 

14  months 

15  months 

Totals 

8  animals 

35  animals 

3  animals 

5  animals 

Reverting  to  the  questions  above  raised,  the  data  show 
that  some  of  the  animals  were  certainly  infested  on 
arrival  here,  and  that  others  probably  were;  but  since 
these  animals  were  not  examined  for  parasites  on  arrival 
in  the  Garden  the  duration  of  infestment  remains 
unknown,  and  accordingly  we  are  not  justified  in  going 
farther  in  our  conclusions.  In  the  case  of  the  marmosets, 
though,  if  we  confine  ourselves  to  the  non-parasitized 
animals,  it  would  appear  that  the  ''acclimatization" 
period  is  within  the  first  six  months.  I  have  attempted 
to  arrive  at  a  conclusion  on  this  basis,  but  the  average 
lifetime  of  the  four  parasitized  marmosets  which  survived 
this  period  is  the  same  as  that  of  the  sixteen  non-para- 
sitized survivors,  and  we  do  not  know  at  what  time  the 
parasitized  ones  contracted  the  disease. 


"^^ 

'  »ii. 


3  ^^ 


y%^ 


\^*' 


Fig.  80. — ARACHNID  (PNELiMUN'i  ^^l.  h  lu\l;  IN  1,U.\(;  OF  ADULT  MONKEY  (MA- 
CACUS  RHESUS).  IT  OCCUPIES  THE  CENTRE  OF  A  CYST  WHICH  IMMEDIATELY 
UNDERLIES  THE  PLEURA  SEEN   AT  UPPERMOST  PART  OF  THE  ILLUSTRATION. 


THE  ANIMAL  PARASITES  647 

Cysticeecus  tenuicollis. — We  have  noted  this  blad- 
der worm  in  the  Aoudad  {Ovis  tragelaphus),  Red  River 
Hog  {Potamochoerus  porcus),  domesticated  Angora 
Goats  and  several  deer  (Cervus  alfredi,  Capreolus  capre- 
olus,  Mazama  mexicana,  M.  hemionus)  located  with  one 
exception  in  the  peritoneal  cavity  or  membrane.  One  of 
the  mule  deer  {Mazama  hemionus)  exhibited  the  parasite 
also  in  the  lung  and  liver.  This  parasite  is  discussed 
because  the  very  valuable  Philippine  spotted  deer 
{Cervus  alfredi)  died  from  a  peritonitis  secondary  to  an 
infected  cyst  in  the  lesser  omentum,  and  because  the 
parasitism  {Tcenia  marginatum)  is  contractible  from 
canidae  which  are  also  on  exhibition  in  the  Garden.  It 
happens  that  the  spotted  deer  did  not  become  infested 
from  the  dogs,  but  it  is  quite  probable  that  the  goats  did, 
since  they  passed  many  times  daily  in  front  of  the  wolf 
cages,  drawing  the  children's  carriages  over  the  walks, 
and  were  stabled  nearby.  We  have  not  discovered  any  of 
the  other  tapeworm  cysts  in  deer  which  might  be  trans- 
mitted to  them  from  the  canidae.  Camels  which  are  parked 
directly  opposite  them  have  only  exhibited  echinocoecus 
cysts,  yet  we  have  never  found  its  adult  form  {Tcenia 
echinocoecus)  or  its  ova  in  the  canine  feces.  The  danger 
of  fatal  disease  from  C.  tenuicollis,  even  though  the  infest- 
ment  be  present,  is  remote ;  but  we  feel  that  it  is  better,  if 
possible,  not  to  exhibit  the  canidae  adjacent  to  sus- 
ceptible animals. 

Pulmonary  Acaeiasis  in  Monkeys. — We  have  seen  but 
two  instances  of  this  affection  in  the  Philadelphia  Garden. 
The  offending  parasite  in  our  animals  was  a  new  species, 
Pneumonyssus  foxi  Weidman  ( 32 ) .  It  occurred  sparingly 
in  small  cysts  under  the  pleura  and  was  certainly  benign 
in  our  cases.  The  importance  of  the  infestment  consists 
in  part  in  that  these  lesions  may  be  mistaken  for  tubercles. 

(32)   Jour.  Parasit.,  Sept.,  1915,  V.  2,  pp.  37-45. 


648  DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

At  the  London  Gardens  (33)  acariasis  was  found  in 
forty-four  young  rhesuses  dying  of  pneumonia,  and  the 
observers  ascribed  the  inflammation  to  irritation  of  cer- 
tain doubly  refractile  crystals  which  occurred  in  the 
excreta  of  the  mite.  There  are  four  other  recorded 
instances  of  such  disease  in  monkeys,  all  caused  by  dif- 
ferent species  of  parasites. 

As  to  pathogenesis  of  these  arachnids,  the  London 
experience  is  most  illuminating  in  that  it  was  young 
rhesuses  that  were  affected.  Our  specimens  were  mature, 
and  nothing  was  stated  to  the  contrary  in  the  other 
reported  cases  from  various  parts  of  the  world.  The 
parasites  are  perhaps  inhaled  from  the  straw  used  as 
bedding,  since  such  vegetable  material  is  a  common 
habitat  for  mites.  If  the  resultant  acute  pneumonia  is 
weathered  the  relics  might  remain  only  in  the  form  of 
the  subpleural  and  parabronchial  cysts  such  as  we  have 
seen  at  the  Philadelphia  Garden. 

I  am  the  more  willing  to  accept  the  possibility  that  the 
simian  arachnids  can  induce  an  acute  pneumonia  after 
studying  a  very  definite  case  of  bronchopneumonia  in  a 
prairie  dog,  which  was  induced  by  Cytoleichus  penrosei 
Weidman  1916.(34) 

Periproventriculae  FiLARro^  or  Birds. — Every  year 
we  report  a  number  of  cases  (up  to  twenty- three)  of  these 
worms,  probably  several  species,  coiled  under  the  serosa 
of  the  air  sacs  and  most  commonly  around  the  proven- 
tricle.  Tentatively  we  have  recognized  two  forms,  a 
shorter  (an  inch  or  so  long)  and  a  longer  (three  to  four 
inches).  The  latter  is  most  inextricably  coiled,  but  the 
former  may  be  teased  out.  Microfilaria  occur  in  the  blood 
of  the  latter  cases,  but  not  in  that  of  the  former.  The 
adults  have  been  observed  to  penetrate  from  their  posi- 
tion in  the  air  sac  serosa  into  the  lumen  of  the  proventricle 
(goose),  to  have  caused  rupture  of  the  inferior  cava 

(33)  Proc.  Zool.  8oc.  Lvndon,   1919,  p.  14. 

(34)  Jour.  Parasit.,  Dec,  1916,  V.  3,  pp.  82-89. 


Fig.    81.— arachnid    (CYTOI.EICHUS   PENROSEI)    IN   A    RRONCHOPNEUMONIC    FOCUS    IN    THE 
LUNG   OF  A  PRAIRIE  DOG    (CYNOMYS  LUDOVICIANUS). 


Fig.   82. — FILARIAL  WORM   COILED   NEAR  PROVENTRICLE 
OF  A   FINCH. 


THE  ANIMAL  PARASITES  649 

(bulbul),  to  be  associated  with  subserous  cysts  of  the 
intestine  (weaver)  and  with  profound  anemia  (liothrix). 
The  birds  affected  are  mostly  small,  inexpensive  ones,  but 
the  infestment  is  important  because  of  its  frequency  and 
deserves  study  of  the  means  of  transmission. 

Physaloptera  in  Opossums  and  Badgers. — These 
worms  were  frequent  findings  for  a  period  of  years  and 
were  particularly  impressive  on  account  of  the  large 
number  of  parasites  present.  The  stomach  often  con- 
tained scores,  more  or  less  securely  attached  to  the 
mucosa  by  the  head.  The  worms  average  an  inch  or  two 
in  length  and  perhaps  an  eighth  of  an  inch  in  thickness. 
P.  turgida  is  the  only  species  we  have  identified  (three 
examinations).  As  to  pathogenicity  we  have  not  observed 
that  definitely  constant  lesions  are  induced  by  the  para- 
sites. In  several  instances  the  gastric  mucosa  has  shown 
the  mosaic  appearance  indicative  of  chronic  gastritis,  a 
condition  not  necessarily  incited  by,  but  certainly  aggra- 
vated by,  these  worms ;  at  least  significant  is  the  habit  of 
the  worm  to  imbed  its  head  in  the  gastric  mucosa.  In  one 
instance  the  microscope  has  revealed  a  most  severe 
fibrosis  of  the  submucosa.  The  fibrosis  was  not  so  much 
diffuse  as  it  was  local  or  nodular,  and  in  favorable  places 
the  ova  of  physaloptera  could  be  discovered  in  the  centres 
of  the  nodules,  and  thus  betrayed  the  previous  presence 
of  the  adult  worm  there.  In  this  individual  animal  the 
case  against  the  physaloptera  is  clinched  by  direct  evi- 
dence. In  other  cases  we  have  circumstantial  evidence. 
Whereas  it  is  not  a.  deeply  burrowing  parasite,  it  is  still  a 
penetrative  one,  and  this  is  sufficient  to  compromise  the 
all  important  * ' integrity  of  the  mucosa, ' '  It  should  there- 
fore be  considered  pathogenic  in  all  cases,  because  open 
to  suspicion  in  several  directions — abstraction  of  tissue 
juices,  irritation  by  its  products  or  movements  and  by 
opening  up  an  avenue  for  bacterial  infection. 

[  Tropidocerca  in  Birds. — This  is  a  blood-red  nematode 
of  the  size  of  a  mustard  seed  to  that  of  a  peppercorn  which 
42 


650   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

inhabits  the  depths  of  the  proventricidar  mucosa.  At  first 
sight  its  spheroidal  form  suggests  that  of  a  fluke,  but 
under  the  microscope  it  is  found  to  be  a  nematode 
hugely  ballooned  out  by  ova,  and  coiled  up  into  a  ball.  In 
spite  of  its  dangerous  appearance — being  red — it  is  most 
likely  quite  innocuous,  for  microscopic  sections  show  no 
sign  of  inflammation  around  the  worm.  Moreover,  we 
know  that  a  Concave  Casqued  Hornbill  /  {Dichoceros 
bicornis)  now  on  exhibition  has  harbored  the  worms,  as 
indicated  by  ova  in  the  droppings,  for  eight  years  and  yet 
seems  perfectly  well.  I  have  made  wax  reconstructions 
of  three  of  the  worms  and  find  that  the  coils  are  not  very 
intricate  and  that  they  assume  no  regular  or  con- 
stant arrangement. 

V  Syngamus  Trachealis. — Our  worst  experience  with 
this  picturesque  parasite  was  in  common  crows  {Corvus 
b.  brachyrhynchos) .  In  1914  and  1915  alone  we  lost  five 
such  birds.  Some  geese,  swans  and  a  pheasant  complete 
the  short  list  of  birds  affected  in  addition  to  the  crows.  In 
no  case  was  it  a  young  bird  that  was  affected.  Shipley  (35) 
reports  this  parasite  in  two  grouse  at  the  London 
Gardens,  and  Plimmer's  tables  show  that  three  deaths 
were  directly  charged  against  them  in  one  year(36). ' 

Extra-intestinal  Tapeworms. — This  discovery  is 
worthy  of  record  because  it  is  rare  for  cestodes  to  appear 
anywhere  save  in  the  intestines.  We  have  observed  three 
instances  where  they  had  backed  up  into  the  bile  duct — 
twice  in  the  Cape  Hyrax  {Procaria  capensis)  and  once  in 
a  Livingston's  Eland  {Taurotragus  oryx  livingstonii). 
At  the  London  Gardens  they  were  mentioned  in  the  gall- 
bladder of  a  wallaby  and  in  Cape  Hyraces.  Beddard(37) 
carefully  describes  four  new  species  of  these  cestodes 
from  the  hyrax. 

(35)  Proc.  Zool.  Soc.  London,  1909,  p.  335. 

(36)  Loc.  cit.,  1912,  p.  236. 

(37)  Loc.  cit.,  1912,  p.  576. 


Fu,.   8?.— PHVSALOPTERA    IN    STOMACH    OK    COMMON     OPOSSLM 
(DIDELPHVS  VTRGIMANA).      THIS    IS    NOT    AN    EXCEPTIONAL     OE(;REE 
OK  INVOLVEMENT. 


i^PMPt' 


l^^M '  ;i: 


r'v 


Fir..  84. — ONE  OF  THE  FIBROUS  NODILES  IN  THE  GASTRIC  Sl'BMlCOS.\  OF  AN 
OPOSSUM.  AN  OVUM  OF  PHYSALOPTERA  IS  SEEN  PRECISELY  IN  THE  MIDDLE  OF 
THIS  ILLUSTRATION. 


THE  ANIMAL  PARASITES 


651 


Table  28. 
Occurrence  of  Blood  Parasites. 
(Adapted  from  Plimmer,  nine  year  period) 
Animals  examined-1 2,241     Mammalia-2,924    Aves-6,619    Reptilia-2,( 


1.  Hemogregarines . 

2.  Microfilaria 


3.  Hemoproteus. . 

4.  Trypanosomes. 


5.  Plasmodia. 


6.  Leucocytozoa 

7.  Intestinal  organisms* 

8.  Toxiplasma 


9.  Spirochaeta 

10.  Babesia 

11.  Haemocystidium . 


Host 


Grand  Total. 


Reptilia 

Mammalia 

Aves 

Reptilia 

Aves 

Mammalia 

Aves 

Reptilia 

Amphibia 

Mammalia 

Aves 

Reptilia 

Aves 

Reptilia 

Mammalia 

Aves 

Reptilia 

Mammalia 

Mammalia 

Reptilia 


No. 
Infested 


316 

33 

191 

24 

140 

1 

28 

4 

3 

2 

39 

5 

16 

16 

1 

1 

1 

1 

1 

1 


824 


%  Infested 


11.8 
1.1 
3. 
1. 
2.1 
0.003 
0.4 


0.6 


0.2 
0.5 


*The  exact  taxonomic  position  could  not  be  stated,— probably  an  ameba. 

Summary  of  Table  28. 


Mammalia. 

Aves 

Reptilia 


Total. 


Parasitized 


39 
415 
367 


821 


Animals 
examined 


2,924 
6,619 
2,698 


12.241 


1.5 

6.5  — 
14.0 


6.7 


FiLARiASis  IN  Wild  Cats  {Felis  ruff  us)  .—This  para- 
site was  named  Filaria  fasciata  because  it  coils  in  the 
fascia  between  the  muscles— generally  those  of  the  thigh 
and  abdomen.  The  worms  are  easily  detected  on  skinning 
the  animal  and  separating  thigh  and  other  muscles. 
Microfilaria  were  always  present  in  the  blood.  The  grade 
of  pathogenicity  is  only  conjectural. 

Peritoneal  Filaria  in  Monkeys.— Thread  worms 
have  been  encountered  eleven  times,  largely  in  Cebidae.  In 


652   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

several  instances  F.  gracilis  has  been  the  species  identi- 
fied, always  inhabiting  the  peritoneal  cavity,  and  in  one 
instance  also  the  lung.  Microfilaria  were  always  present 
in  the  blood.  We  have  never  seen  lymphangitis  or 
elephantiasis  in  our  filarial  cases. 

Blood  Parasites. — I  justify  this  paragraph  on  the 
basis  of  the  usefulnes  it  might  have  in  the  clinical  direc- 
tion, for  while  the  taking  of  blood  specimens  is  not  as 
easy  as  with  man  it  can  still  be  done  wdth  some  animals. 
From  time  to  time  we  have  encountered  blood  parasites 
in  this  Garden,  but  the  large  numbers  occurring  in  the 
experience  of  special  searchers  in  the  London  Garden 
and  Plimmer  's  particular  interest  in  this  direction  make 
their  data  much  the  more  valuable.  In  one  report  of  6,430 
animals  examined  he  found  7  per  cent,  infested  with  blood 
parasites  of  one  sort  or  another.  I  have  constructed  the 
foregoing  table  (28)  from  his  various  reports  to  show 
which  animal  classes  were  affected  by  the  several 
blood  parasites. 

This  table  (28)  brings  out  that  considering  them  as  a 
whole  and  without  respect  to  host,  just  as  the  animals 
come  day  in  and  day  out  to  the  autopsy  table,  blood 
parasites  will  be  met  in  6.7  per  cent,  of  all  cases.  They 
are  seen  most  commonly  in  the  form  of  hemogregarines 
of  reptiles  (2.5  per  cent,  of  all  animals  and  12  per  cent, 
of  all  reptiles)  while  microfilaria  run  a  close  second,  being 
found  in  2  per  cent,  of  all  animals  but  much  more 
commonly  in  birds.  Hemoproteus  of  birds  while  ranking 
third,  should  be  emphasized  on  account  of  its  acknowl- 
edged blood-destructive  properties.  The  remaining  infes- 
tations were  too  infrequent  to  be  useful  statistically. 

Turning  to  individual  groups  of  (blood  parasites,  micro- 
filariaB  of  birds  deserve  special  comment.  They  occurred 
four  times  more  often  in  birds  than  in  other  animals,  or, 
put  in  another  way,  one  out  of  every  twenty-two  birds 
was  affected,^  and  only  one  out  of  every  ninety  other 
animals.  (  The  high  figure  for  birds  is  significant  in  rela- 


Fig.  85. — AD.'VPTATION  FROM  RECONSTRL'Cl  l()\  ol  IROPl  IK)CERC.\  C 
THE  WORM  LAY  IN  THE  WALL  OF  THE  PROVENTRICLE  OF  A  LOUISIANA  HER( 
TRICOLOR   RUFICOLLIS). 


MM.     . 
1 

ijii  nil 

1  2 

iltiiiil. 

nlilifH 

,,1 


bl 


(iiilniiliiimiitliinliiitltiiiHit^ 


86.— CESTODES   (THREE)   PROJECTING   FROM  THE  SEVERED   END   OK  THE 
DUCT  OF  A  CAPE  HYRAX   (PROCARIA  CAPENSIS). 


THE  ANIMAL  PARASITES  653 

tion  to  what  we  have  already  said  about  periproven- 
tricular  filaridae  in  our  Garden,  indicating  that  the  same 
inf  estment  probably  also  exists  in  London. 

A  point  brought  out  by  Plimmer  is  to  the  effect  that, 
of  the  several  blood  parasites,  the  microfilariae  were  the 
least  harmful,  and  that  of  these  the  adult  forms  were  the 
only  ones  to  produce  symptoms;  yet  in  one  place (38)  he 
records  microfilaria  as  plugging  the  cerebral  capillaries 
of  birds.  This  is  a  very  important  lesion  if  permanent, 
and  especially  so  when  affecting  cerebral  capillaries  as 
do  the  organisms  and  pigment  of  malaria.  The  adult 
forms  were  found  in  one-fourth  of  the  cases  where  micro- 
filaria were  demonstrated. 

As  to  the  pathogenicity  of  these  blood  parasites  in 
general,  it  will  be  unsafe  to  arrive  at  a  definite  conclusion, 
recalling  the  pitfalls  that  I  have  already  outlined  in  dis- 
cussing pathogenicity  of  parasites  in  general.  Keeping 
in  mind  the  wonderful  adaptability  on  occasion  of  animals 
to  unfavorable  circumstances  we  must  hesitate  to  declare 
unqualifiedly  the  importance  of  even  blood  parasites  as 
morbid  agents.  Where  the  parasite  is  known  to  destroy 
the  blood  cells  of  birds  and  mammals  it  is  otherwise,  but 
even  here  experimental  work  would  be  necessary  to  settle 
the  question.  The  element  of  ' '  racial  ' '  immunity  and  of 
phylogeny  is  the  fly  in  the  ointment  of  our  deductions.) 

Transmission  of  Animal  Parasitism  From  Wild  Ani- 
mals TO  Man. 

Examples  of  direct  transmission  will  be  only  occa- 
sional, due  to  the  relatively  infrequent  contacts  between 
the  two  hosts.  Pets  threaten  the  most.  Several  such 
examples  have  been  touched  upon  in  the  preceding  pages 
and  it  but  remains  to  gather  them  into  one  place.  There 
is  one  concrete  instance  in  the  form  of  clear-cut  simian 
scabies  being  transmitted  to  a  keeper  in  this  Garden (39) 

(38)  Proc.  Zool.  Soc.  London,  1910,  p.  134. 

( 39 )  Weidman  ( F.  D. ) ,  "  Dermatoses  of  Monkeys,"  Arch.  Derm,  and 
Syph.,  Chicago,  March,  1923,  p.  289. 


654   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

and  a  similar  lot  fell  to  the  keeper  of  a  wombat  at  the 
Paris  Garden  (40)  as  well  as  to  the  taxidermist  who  pre- 
serv'ed  its  skin.  We  know  that  the  skin  and  feathers  of 
our  parrots  and  pigeons  harbor  mites (41)  (plumicoles 
of  Megnin)  and,  recalling  the  occasional  cases  of  poultry- 
men's  itch,  a  transient  affection  might  be  conceded  from 
pet  parrots  and  other  birds.  Pediculi  are  not  as  num- 
erous on  monkeys  as  popularly  supposed — ^we  see  very 
few  at  the  autopsy  table.  We  have  seen  Trichinella  spira- 
lis in  the  polar  bear  {Ursus  maritimus) — an  animal  whose 
flesh  is  edible.  The  hydatid  cysts  in  the  camel  appear  un- 
important, but  in  the  livers  of  deer  it  is  otherwise. 
Neither  of  these  infestments  is  dangerous  if  the  meat  is 
sufficiently  cooked  before  eating. 

Hookworm  disease  points  thus  far  only  to  Anchylos- 
toma  duodencde  in  the  chimpanzee  and  Uncinaria  cey- 
lanicum  in  the  lion  and  tiger.  Both  serve  as  reservoirs 
of  the  disease,  the  ova  being  discharged  by  way  of  the 
feces.  Similarly  the  Strongyloides  intestinalis  infest- 
ment  which  we  have  seen  in  the  orang  might  be  trans- 
ferred to  man.  Indirectly,  Europeans  traveling  in  Africa 
have  made  the  crucial  test  that  certain  ungulates  and 
other  wild  animals  of  Africa  are  the  reservoirs  of  Try- 
panosoma gamhiense,  the  parasite  of  the  well  known 
African  sleeping-siclmess ;  for  this  example  the  blood 
stream  of  the  beast  is  the  reserv^oir  and  a  biting  insect 
the  means  of  transmission. 

The  above  examples  are  cited  to  emphasize  the  pos- 
sibility that  parasites  of  wild  animals  may  have  a  patho- 
genic significance  for  man.  They  do  not  exhaust  the 
subject.  Many  more  instances  might  be  cited  but  the  fore- 
going bring  out  the  important  ones  which  have  come  to 
our  attention. 

(40)  Railliet  Traite  de  Zool.  Med.  et  Agric.  Paris — Asselin  et 
Houzeau,  1895,  p.  659. 

(41)  Megnin,  Les  Parasites  Articules,   1895,  Masson  et  Cie,  Paris. 


THE  ANIMAL  PARASITES  655 

Treatment. 

The  recognition  of  the  existence  of  parasites  during 
the  life  of  an  animal,  especially  those  of  the  sldn  and 
intestinal  tract  whose  discovery  is  easiest,  suggests  that 
some  means  of  combating  them  should  be  employed.  But 
we  are  by  now  quite  satisfied  that  medicinal  and  disin- 
fective  therapeutic  procedures,  while  they  ha\^e  their 
field  of  usefulness,  are  much  less  to  be  depended  upon 
for  the  protection  of  exhibits  than  are  preventive  meas- 
ures of  general  hygienic  nature.  Under  the  latter  head- 
ing come  the  prompt  removal  of  excreta,  frequent  changes 
of  drinking  water,  routine  examinations  of  feces  of  cer- 
tain varieties,  autopsy  examinations  and  incineration  of 
autopsy  remains — all  of  which  are  part  of  the  require- 
ments of  common  cleanliness  and  general  disease  preven- 
tion. I  wish  to  amplify  the  matter  of  disposal  of  feces 
and  general  cage-police.  Our  ideas  as  to  what  constitutes 
thoroughness  in  this  work  have  changed  considerably 
since  Fulleborn's  recent  demonstration  that  ascarid  ova 
(42)  could  live  in  formaldehj^de  for  four  or  five  years, 
and  the  older  one  of  Galli-Valerio  (43)  that  those  of 
Hepaticola  hepatica  lived  one  month  in  2  per  cent,  formal- 
dehyde solution.  Evidently  the  same  substances  which 
disinfect  do  not  invariably  disinf est ;  and  if  the  occasion 
should  arise  for  the  most  exacting  control  in  tliis  respect, 
a  special  investigation  of  the  susceptibility  of  the  indi- 
^4dual  ova  in  question  would  have  to  be  undertaken. 

In  addition  to  these  general  measures  we  have  put 
up  certain  special  safeguards  against  parasites.  Thus, 
each  specimen  of  the  large  Carnivora  (lions,  tigers,  leop- 
ards, etc.),  has  received  routinely  a  dose  of  santonin 
every  month  over  a  period  of  several  years.  We  have 
no  figures  on  which  to  base  comparison  with  previous 
periods,  but  an  examination  of  feces  of  all  the  inmates 

(42)  Quoted  by  Jensen  (V.),  Hospitalstidende,  Copenhagen,  1922,  65, 
No.  28.  p.  457. 

(43)  Centr.  f.  Bakt.  u.  Parasitk.,  (etc.),  Jena  1— Abt.  V.  35  (1), 
5,  1903,  orig.  p.  89. 


656   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

of  the  Camivora  house  in  1916(44)  showed  that  less  than 
one-third  of  the  animals  were  infested,  and  of  these 
all  save  the  jaguars  showed  either  small  numbers  of  ova 
in  the  feces  or  relatively  non-pathogenic  forms.  The 
jaguars  had  been  badly  infested  for  many  years  with 
dibothriocephalus.  Prior  to  this  examination  we  had 
been  under  the  impression  that  nearly  every  one  of  the 
felidae  ordinarily  was  infested  and  if  this  impression 
was  well  founded,  due  credit  must  be  given,  in  company 
with  general  hygienic  precautions,  to  the  routine  santo- 
nin dosages.  It  goes  without  saying  that  where  animals 
are  detected  at  autopsy  with  unequivocal  transmissible 
and  dangerous  parasites  (coccidia,  amebae,  etc.),  the  con- 
tacts are  isolated,  examined  and  if  necessary  treated  for 
the  affection  or  even  sacrificed. 

To  continue  the  preventive  measures,  it  would  be  most 
desirable  to  examine  at  least  the  blood  and  feces  of  all 
newly  arrived  animals,  but  at  present  this  is  not  practi- 
cable on  account  of  the  labor  involved  in  the  laboratory 
and  in  collecting  the  material,  and  because  all  animals 
do  not  stand  the  restraint  involved  when  blood  specimens 
are  being  taken.  At  present  we  are  limiting  special  ex- 
aminations to  the  droppings  of  newly  arrived  parrots  and 
toucans  for  Spiroptera  incerta  and  to  the  feces  of  certain 
monkeys  for  amebae. 

Further  preventive  measures  will  depend  on  the 
nature  of  individual  infestments  as  they  crop  up.  Food 
inspection,  screening,  sulphur  dips,  etc.,  are  but  a  few 
examples  of  what  might  be  found  necessary  hygienically 
after  investigating  or  establishing  the  life  cycle  of  our 
numerous  parasitic  groups.  However  we  cannot  forbear 
to  emphasize  again  the  value  of  the  blast  lamp  and  of 
paint  in  the  hygiene  of  animal  enclosures — ^means  we 
believe  to  be  much  more  potent  and  quite  as  practicable  as 
chemical  disinfectants. 

(44)    Phila.  Zool.  Soc.  Rep.,  1917,  p.  36. 


Fig.  87. — TRICHINELLA  SPIRALIS  IN  MUSCLES  OF  POLAR  BEAR  (URSUS  MARI- 
TIMUS).  THIS  WAS  AN  OLD  INFESTMENT.  AS  INDICATED  BY  THE  THICK  AND  HYA- 
LOID  CHARACTER   OF  THE  CAPSULE. 


THE  ANIMAL  PARASITES  657 

Turning  now  to  the  active  curative  side  of  the  sub- 
ject, what  medical  means  we  have  against  parasites  ap- 
pertain for  the  most  part  to  the  intestinal  ones.  The 
treatment  of  tapeworms  is  very  hazy  and  unsatisfactory 
— areca  nut  is  perhaps  more  useful  in  animals  than  any 
one  other  drug.  For  round  worms  santonin  is  most  to 
be  depended  on  although  turpentine  is  useful  against  the 
round  worm  of  the  Equidae.  The  dosage  of  santonin  per 
month  has  been— for  large  bears,  ten  grains ;  for  lions, 
tigers,  large  pumas,  six  grains;  for  jaguars,  leopards, 
hyenas,  four  grains ;  for  wild  cats,  etc.,  two  grains.  The 
dose  of  areca  nut  recommended  for  Carnivora  is  two 
grains  per  pound  of  body  weight.  Since  ung-ulates  do  not 
stand  areca  nut  well,  iron  sulphate  may  be  used.  For 
animals  the  size  of  a  horse  the  dosage  is  two  drams,  and 
to  this  one  or  two  grains  of  arsenic  trioxide  may  be  added. 
On  the  basis  of  very  carefully  controlled  experiments  on 
dogs.  Hall  recommends  carbon  tetrachloride  for  hook- 
worms in  these  animals — 0.3  mils  per  kilo  of  body  weight, 
without  purging.  Its  efficacy  has  been  confirmed  lately 
but  we  have  not  had  the  occasion  to  test  it. 

Prom  time  to  time  we  have  broached  other  lines  of 
medication  against  worms  which  may  be  worth  while  re- 
lating if  for  nothing  more  than  to  illustrate  the  uncertain 
ways  of  our  vermifuges  when  applied  to  wild  animals. 

I  can  speak  first  of  thymol  as  employed  on  parrots 
parasitized  by  Spiroptera  incerta.  The  first  thing  that 
impressed  us  was  the  large  dosage  which  birds  could  en- 
dure. The  lethal  dose  for  pigeons  was  four  grains,  sus- 
pended in  mucilage  of  acacia.  After  we  had  established 
that  certain  parrots  withstood  fourteen  grains  in  muci- 
lage, we  administered  on  one  occasion  twelve  grains  and 
on  another  sixteen  grains,  suspended  in  glycerin.  The 
drug  is  reputed  to  be  absorbed  when  exhibited  in  the 
latter  vehicle  and  we  hoped  to  get  a  certain  anthelmintic 
effect  on  the  parasites  from  the  blood  side  as  well  as 
from  the  lumen  of  the  gut.    The  bird  itself,  a  very  heavily 


658   DISEASE  IN  WILD  MAMMALS  AND  BIRDS 

infested  cockatoo,  showed  no  ill  effects  and  passed  two 
dead  female  spiroptera  and  enormous  numbers  of  ova. 
But  thereafter  it  passed  even  greater  numbers  of  ova 
than  before  (we  estimated  182,000  per  day  for  this  bird 
over  a  five  day  period  and  288,000  on  a  single  subsequent 
day),  and  was  obviously  unimproved  by  the  treatment. 
The  explanation  of  failure  was  clear,  for  the  worms  can 
retire  into  the  protecting  mucus  or  mucous  membrane 
lining  the  proventricle  until  the  thymol  has  passed  by, 
and  even  though  paralysed  may  not  be  flushed  out.  In 
a  later  test  on  a  toucan  which  died  twenty  minutes 
after  thymol  administration  we  found  at  the  autopsy 
that  worms  deeply  imbedded  in  the  proventricle  were 
translucent  and  motionless  from  the  effects  of  the 
thymol-glycerin  mixture,  i.e.,  saturated  with  the  medi- 
cament and  apparently  dead.  Twenty  minutes  later 
they  were  placed  in  normal  salt  solution  in  the  incu- 
bator, and  next  morning  were  found  actively  motile. 
Thymol  evidently  does  not  kill — it  only  stupefies,  and  in 
the  absence  of  means  for  flusliing  the  parasites  out,  as  we 
do  in  human  hookworm  cases,  this  class  of  vermifuge  will 
have  to  be  abandoned  in  work  against  tliis  parasite. 

Not  with  any  serious  hope  of  success,  but  feeling  that 
arsenic  was  the  most  promising  drug  available  for 
parenteral  use,  we  tried  atoxyl  hypodermically  and 
arsphenamine  intravenously  but  without  success.  The 
only  positive  results  were  to  emphasize  the  tolerance  of 
some  lower  animals  to  arsenic.  Thus  in  preliminaiy  work 
pigeons  received  sixty  drops  of  Fowler's  solution  by 
mouth  without  embarrassment,  but  five  minims  killed  a 
pigeon  when  administered  hypodermically.  The  organic 
arsenical,  arsphenamine,  w^as  withstood  intravenously  by 
pigeons  in  six  times  the  proportional  human  dosage. 

One  of  our  drug  trials  was  instructive  in  that  it 
worked  quite  a  different  effect  from  that  in  man,  besides 
being  most  amusing.  In  earlier  diagnostic  work  on 
spiroptera  we  tested  the  practicability  of  examining  the 


THE  ANIMAL  PARASITES  659 

vomitus  for  the  worms,  hoping  thereby  to  get  a  greater 
concentration  of  ova,  which  would  facilitate  the  micro- 
scopic examination.  Hypodermic  injections  of  apomor- 
phine  (0.1  grain)  into  an  amazon  did  not  induce  vomiting 
from  the  gizzard  as  hoped — only  a  regurgitation  from  the 
crop,  but  it  did  cause  some  dizziness  and  most  ludicrous 
talking  and  laughter. 

To  illustrate  further  the  difficulties  of  animal  medi- 
cation I  quote  our  experience  with  four  red  howling 
monkeys  {Alonatta  seniculus).  One  of  these  died  of  intes- 
tinal obstruction  from  large  ascarids — the  case  which 
has  been  already  cited.  Ova  were  found  in  the  stools  of 
the  remaining  three,  and  one  of  the  monkeys  was  treated 
twice  mth  santonin.  It  died  in  thirty  hours  after  the 
second  dose — not  of  santonin  poisoning,  for  none  of  the 
clinical  symptoms  were  present,  but  most  likely  from  ab- 
sorption of  toxic  substances  originating  in  the  decom- 
posing ascarids  wliich  crowded  the  gut.  It  profits  not 
to  destroy  these  parasites,  then,  unless  we  feel  assured 
that  they  may  thereafter  be  removed  immediately. 

If,  for  the  sake  of  brevity,  I  were  asked  to  state  in  a 
single  sentence  the  practical  status  of  animal  parasitic 
disease  in  this  Zoological  Garden  I  would  put  it  thus: 
Since  there  are  various  animal  parasitic  diseases  con- 
tinuously present  here  of  which  we  know,  and  since  fresh 
ones  are  from  time  to  time  cropping  out,  and  since  these 
are  on  the  whole  of  economic  importance,  it  behooves  us 
to  continue  and  extend  our  efforts  against  an  issue  extant 
— somewhat  through  therapeutic  means,  but  far  more 
through  clinical  laboratory  examinations,  careful  au- 
topsy searches,  and  by  rigid  general  hygienic  measures 
such  as  cage-police,  new  quarters,  isolation,  or  if  neces- 
sary, destruction  of  the  exhibit. 


PROPf 

N.  C, . 


INDEX 


Abortion,  305 
Abscess  of  liver,  231 

of  lung,  155 
Acariasis,  lungs,  647 
of  monkeys,  647 
Actinomycosis,  138,  568 
in  deer,  368,  568 

tapirs,     568 
treatment,    570 
Adenoma,  474 
Adrenal  body,  336 
Alimentary    tract,    166 
Amblyopia,    403 
Amoebae,     606, 644 

dysentery  from,  644 
Amyloid,    liver,   227 

spleen,     128 
Anatomy  of  labor,  290 
Anchylostomum,  see  hookworms 
Anemia,    87 

primary,     98 
secondary,  88 
Aneurysms,  65,  80 
Animal  Parasitism,  hygiene,  656 
prevention,  656 
treatment,  655 
Animal  Parasites,  614 

disappearance  of,  627 
frequency,  628 

of  groups,  633 
in  blood,  652 
incidence,  628-636 
modes  of  action,  617 
occurrence  in  wild,  627 
transmission  animals  to  man, 

653 
visceral  distribution,  637 
Angina  pectoris,  49 
Aorta,  72 

fatty  deposits  in,  71 
Arteries,  66 
Arteriosclerosis,  71 
Arteritis,  70 
Arthritis,  347 

gouty,  347,  411 
Ascending  nephritis,  276 
Aspergillosis,  558 
Aspergillus,  varieties,  558 
Ataxia,  375 

Atrophy,  acute  of  liver,  228 
Autopsy  Ust,  47 
Avian  spiropteriasis,  172,  640 


Bacterial  flora,  418 
Basal  cell  carcinoma,  475 
Beriberi,  439 
Bihary  tract,  225 

calculi,  238 
Birth  canal,  287,  296 

comparative  anatomy,  287  et  seq 

obstructions  to,  306 
Blackhead,  206 
Bladder,  gaU,  224,  238,  239 

urinary,  286 
Blood,  diseases  of,  83 
Blood  formation  in  birds,  98 
Blood  vessels,  66 
Bone  marrow,  83,  109,  111 
Bones,  diseases  of,  343 

effects  of  trauma,  343 

tumors  of,  368 
Botryomycosis,  564,  602 
Botuhsm,  604 
Brain,  385 

tuberculosis  of,  378 

tumors  of,  384 

weight  of,  385 
references  to,  387 
Breast,  312 
Bronchi,  141 
Bronchiectasis,  144 

Cage  palsy,  349 
CalcuU,  bihary,  238 

renal,  282 
Carcinoma,  476 

basal  cell,  475 
Cataract,  403 
Cecum,  211 
Cestodes.  637 
Cholangitis,  239, 256 
Cholecystitis,  239 
Choledochitis,  239 
Cholehthiasis,  238 
Chondroma,  472 
Cloaca,  211 
Coccidiosis,  606 
Cirrhosis  of  hver,  232 
Comparative  anatomy  of  uterus,  287 

of  pelvis,  297-303 
Conjunctivitis,  402 
Constipation,  209 
Constitutional  diseases,  410 
Convulsions,  373 
Cornea,  403 
Coronary  arteries,  49 
Cowper's  gland,  313 

661 


662 


INDEX 


Cretinism,  320.  331 
Cvsticercus  tenuicollis,  647 
Cystitis,  286 
Cytoleichus  penrosei,  647 

Deficiency  diseases,  438-443 
Degenerations  of  kidney,  269 

of  liver,  228 
Diabetes,  412 
Diet,  carnivorous,  452 

herbivorous,  452 
grain,  455 
seed,  454 
soft,  453 

omnivorous,  402 

relation  to  disease,  415 
alimentary  tract,  417 
Dilatation  of  heart,  54 
Diphtheria,  600 
Dislocations,  345 
Distemper,  599 
Diverticula  of  intestine,  219 
Diverticulitis,  219 

Dysentery,  amoebic  in  monkevs,  644 
Dystocia,  292 

Ear.  409 

Echinococcus,  647 
Emphysema,  161 
Encephalomyelitis,  380 
Endocarditis,  52 
Endometritis,  305 
Endothehoma,  165,  474 
Enteritis,  177 

in  Aves,  202,  205 
Mammalia,  185 
Enterohepatitis,  605 
EpitheUoma,  475 
Esophagus,  169 

Exophthalmic  goitre,  320,  323,  329 
Eye,  402 

tuberculosis  of,  402 

Fallopian  tubes,  305 
Fat  infiltrations  of  kidney,  268 
hver,  226 
metabolism,  445 
Fibroma,  472 
Filaria,  fasciata,  651 

gracilis  in  monkeys,  651 
in  blood,  652 
fascia,  651 
muscles,  651 
wildcats,  651 
periproventricular,  648 
peritoneum,  651 
Food,  415 

definition,  415 

in  relation  to  alimentary  tract, 
417 


Food,  disease,  422 

poisoning,  457 
Fowl  cholera,  598 

plague,  598 

typhoid,  598 
Fractures,  344 

Gall  stones,  238 

Gas  bacillus  infection,  602 

Gastritis.  204 

GastroenterocoUtis  in  Ungulata,  194 

in    MarsupiaUa,  198 
Gangrene  of  lung,  155 
Giraffe,  hookworm  in,  644 
Gout,  53,  410 

Heart,  dilatation  of,  54 

hypertrophy  of,  54 

diseases  of,  48 

effects  of,  55 

effect  of  strain,  55-59 

weight  of,  63 

relative  vulnerabihty  of,  61 
Hemorrhagic  septicemia,  598 
Hemoglobinuric  fever,  603 
Hemorrhoids,  218 
Hepaticola  hepatica,  641 
Hepatitis,  228 
Hernia,  216 

Heterakis  in  avian  ceca,  606 
Hookworms,  643,  654 

in  giraffe,  644 
Hypernephroma,  339,  341,  342,  475 
Hypertrophic  periosteitis,  346 
Hyperthyroidism,  320 
Hypertrophy  of  heart,  54 

in  Aves,  60 
Hypothyroidism,  320 

Ileus,  213,  261 
Infantihsm,  433 
Infiltrations  of  kidney,  268 

liver,  226 
Inorganic  salts  in  diet,  427 
Intestinal  obstruction,  212 
tract,  177 

inflammation  of,  181 
mechanical  obstruction  of ,  212, 

617 
relation  to  food,  422 
Intestines,  diverticula,  219 

tumors  of,  220 
IridocycUtis,  402 

Kangaroo  disease,  570 

bacteriology,  576,  586 
course  of  attack,  573 
pathology,  575 
prevention,  572 
treatment,  591 


r 


INDEX 


663 


\ 


Kidney,  263 

abscess,  268,  278 
calculi,  282 
degenerations  of,  269 
hemorrhages,  271 
hypertrophy  of,  267 
infiltrations  of,  268 
tumors,  284 
weight  of,  265 

Labor    from    a    comparative    stand- 
point, 290 

obstructions  to,  306 
Laryngitis,  139 
Larynx,  138 

Leontiasis  ossium,  359,  472 
Leucemia,  104 

in  birds,  108 

lymphatic,  105 

myeloid,  109 
Leucocytes,  84-86 
Limberneck  of  ducks,  604 
Lipoma,  472 
Liver,  222 

abscess,  231 

acute  atrophy,  228 

amyloid,  227 

cirrhosis,  232 

degenerations,  228 

fatty  changes,  226 
infiltration,  226 

inflammation,  228 
chronic,  232 

necrosis  in,  230 

tumors,  240 
Lungs,  146 

abscess,  155 

congestion,  148 

gangrene,  155 

infarct,  160 

tumors  of,  162 
Lymphadenitis,  117 
Lymphatic  leucemia,  105 

tissue,  114 

hyperplasia  of,  115 
in  pharyngeal  wall,  115,  138 
Lymph  nodes,  114 

tuberculosis  of,  121 
tumors  of,  122 
Lymphomatosis,  118 

Malnutrition,  424 
Mammary  gland,  312 
Marmosets,  parasites  of,  645 
Marrow  of  bone,  83,  109,  111 
Meningitis,  376 
Metabolism,  carbohydrate,  443 

fat.  445 

inorganic,  427 

protein,  447 


Miliary  tubercle,  avian,  512 
bovine,  510 
human,  511 
monkey,  511 
Miscarriage,  305 
Molluscum  contagiosum,  601 
Monckeberg  sclerosis,  74,  76 
Monkey's  temperature,  520-528 
Moon  blindness,  405 
Muscles,  370 
Mycosis,  137,  558 

of  esophagus,  168 

histology  of,  561 

hygiene,  563 

incidence,  562 

of  lung,  562 

method  of  action,  560 

pharynx,  168,  564 

types  of,  560 
Myelitis,  350,  381 
Myeloma,  111 
Myocarditis,  52 
Myocardium,  49,  50,  65 
Myxcedema,  320,  331 

Necrosis,  liver,  230 

spleen,  130 
Nematodes,  636 
Neoplasms,  462 

incidence  of,  463,  468 

embryonic  origin,  471 

in  captivity,  469 

in  the  wild,  462,  476 

metastasis,  471 

visceral  origin,  477 
Nephritis,  271 

ascending,  276 

effects  of,  280 

histology  of,  279 

toxic,  275 
Nervous  system,  372 
Nocardia  macropodidarum,  585 
Nocardiosis,  570 

Obesity,  446 

Ophthalmia,  periodic,  405 
Osteitis,  346 

Osteitis  deformans,  359,  431 
Osteoma,  368 
Osteomalacia,  349 
Ovary,  cysts,  307 

Pachymeningitis,  externa,  331,  377 
Paget's  disease,  359,  431 
Pancreas,  244 

degenerations,  250 

tumors,  259 
Pancreatitis,  250 

Parasites,  see  animal  parasites,  614 
Parovarian  cyst,  307 


664 


INDEX 


Pasteurelloses,  597 

Pearl  disease,  491,  501,  505 

Pellagra,  441 

Pelvis,  comparative  anatomy,  297-303 

Penis,  313 

Pericarditis,  53 

Pericardium,  position  of  effusion  in, 

54 
Perio&teitis,  hypertrophic,  346 
Periproventricular  worms,  648 
Perisplenitis,  131 
Peritoneum,  260 

tumors,  262 
Peritonitis,  260 
Pharyngitis,  168 
Pharynx,  168 
Phimosis,  313 
Physaloptera  turgida,  649 

in  opossums,  649 
Plants,  poisonous,  459 
Pleura,  163 
Pleuritis,  164 
Pneumonia,  149 

broncho,  152 

fibrinous,  151,  153 
in  Aves,  153 
origins  of,  154 

lobar,  151 
Pneumonokoniosis,  159 
Pneumonyssus  foxi,  647 
Poisonous  plants,  459 
PoliomyeUtis,  380 
Prostate  gland,  313 

enlargements  of,  314 
tuberculosis  of,  315 
tumors  of,  314 
Proventricle,  171 

worms  in,  172,  640 
Psittacosis,  208,  597 
Pyelonephritis,  277 

Quail  disease,  608 

Rabies,  602 
Rachitis,  349,  429 
Rectum,  prolapse  of,  218 
Reproductive  organs,  female,  287 

male,  317 
Respiratory  tract,  134 
Rhinitis,  135 
Rickets,  349,  429 
Renal  calculi,  282 

Salpingitis  fallopii,  305 
Santonin,  657 
Sarcoma,  471,  474 
Scurvy,  440 
Seminal  Vesicles,  315 
Sinusitis,  135 
Skeleton,  343 


Spinal  cord,  373 
Spiroptera  incerta,  638,  640 
detection,  640 
eradication,  640 

in  parrots,  172,  208,  640 
Spiropteriasis,  172,  640 
Spleen,  114,  122 

amyloid,  128 

congestions,  125 

enlargements,  124 

hemorrhage,  125 

inflammation,  126 

in  anemia,  130 

in  hepatic  cirrhosis,  130 

necrosis,  130 

size,  124 

tuberculosis  of,  132 
Squirrel  monkeys,  parasites  of,   645 
Starvation,  425 
Stomach,  174 

tumors  of,  176 

ulcers  of,  175 
Streptothricosis,  567 
Suprarenal  body,  336 
Syngamus  trachealis,  140,  650 

Taenia  echinococcus,  647 
Tape  worms,  637 

in  Uver,  650 
Temperature  of  monkeys,  520-528 
Testes,  313 

tumors  of,  313 
Tetanus,  602 
Thrombosis,  69 
Thymol,  657 
Thymus,  120,  336 
Thyroid  body,  316 

atrophy  of,  330 
hyperplasia  of,  325 
size  of,  318 
tumors  of,  333 
Tonsils,  115,  138 
Trachea,  140 

Tropidocerca  contorta,   649 
Tubercle  bacillus,  types  of,  513 
Tuberculin  test  on  monkeys,  518 
other  animals,  549 
dose,  529 

effect  on  kidneys,  548 
eye,  546 
reaction,  530 
skin,  546 
Tuberculoma,  505 
Tuberculosis  of  brain,  378 

avian  characters,  503,  512 
Carnivora,  498 
control,  514-548 
diagnosis  of,  514 
discovery  during  life,  514 
distribution  in  birds,  504 


91 


INDEX 


665 


Tuberculosis  of  eye,  402 
gelatinous,  504 
histology,  510 
hygiene,  516 
in  Aves,  503 
in  MammaUa,  492 
in  Primates,  492 
in  various  avian  orders  506-510 
incidence,  489 
intestinal  in  birds,  505 
Lemures,  495 
lymph  nodes,  121,494 
nonsusceptible  animals,  490 
ordinate  characters,  492 

frequency,  489 
pathological  type,  490 
Proboscidea.  502 
Rodentia,  499 
routes  of  infection,  485 


Tuberculosis,  sanitation  of  cages,  516 
susceptible  animals,  490,  515-517 
Ungulata,  500 
visceral  distribution,  491 

Tumors,  see  neoplasms 

Ulcer,  gastric,  175 

Uncinaria,  643 

Uremia,  281 

Urethra,  315 

Uterus,  comparative  anatomy,  287 

inflammation,  305 

tumors  of,  308 

Vitamins,  438 
Waterfowl  epizootic,  604 
Zoological  list,  43 


A    LIST    OF    THE    PUBLICATIONS    FROM    THE 

LABORATORY    OF    COMPARATIVE    PATHOLOGY 

OF  THE  PHILADELPHIA    ZOOLOGICAL  SOCIETY 

1909—1923 

1.  Results  of  Tuberculin  Tests  in  Monkeys  at  the  Philadelphia 

Zoological  Garden,  by  C.  Y.  White,  M.D.  and  Herbert 
Fox,  M.D.  The  Archives  of  Internal  Medicine,  December, 
1909,  Vol.  4,  pp.  517-527,  Chicago,  Illinois. 

2.  Note  on  the  Occurrence  of  a  Ciliate  {Opalinopsis  nucleolobata, 

n.s.)  in  the  Liver  of  a  Mammal  (Canis  latrans),  by  Allen 
J.  Smith,  M.D.  and  Herbert  Fox,  M.D.  University  of 
Pennsylvania  Medical  Bulletin,  February,  1909,  Philadel- 
phia, Pennsylvania. 

3.  The  Tuberculin  Test  in  Monkeys:  with  Notes  on  the  Temper- 

ature of  Mammals,  by  Arthiur  Erwin  Brown,  D.Sc. 
C.M.Z.S.,  Sec.  Zool.  Soc,  Phila.  Proceedings  of  the  Zoo- 
logical Society  of  London,  1909,  pp.  81-90. 

4.  Observations  on  the  Occurence  of  Neoplasms  in  Wild  Animals, 

by  C.  Y.  White,  M.D.  and  Herbert  Fox,  M.D.  Proceedings 
of  the  Pathological  Society  of  Philadelphia,  February,  19 10. 

5.  Observations  on  the  Comparative  Anatomy  of  the  Female 

Genitalia,  by  Edward  A.  Schtimann,  M.D.  American 
Journal  of  Obstetrics  and  Diseases  of  Women  and  Children, 
Vol.  LXIV,  No.. 4,  191 1,  New  York. 

6.  Observations  Upon  Neoplasms  in  Wild  Animals  in  the  Phila- 

delphia Zoological  Garden,  by  Herbert  Fox,  M.D.  The 
Journal  of  Pathology  and  Bacteriology,  Vol.  XVII.  (191 2), 
pp.  217-231.     England. 

7.  A  Study  of  Metazoan  Parasites  Found  in  the  Philadelphia 

Zoological  Garden,  by  Fred  D.  Weidman,  M.D.  Proceed- 
ings of  the  Academy  of  Natural  Sciences  of  Philadelphia , 
March.  19 13,  pp.  126  to  151,  Philadelphia,  Penna. 

8.  The  Pathology  of  the  Thyroid  Gland  in  Wild  Animals,  by 

Herbert  Fox,  M.D.  Journal  of  Comparative  Pathology  and 
Therapeutics,  Vol.  27,  p.  23.     Edinburgh,  Scotland. 


A  LIST  OF  PUBLICATIONS 

9.  The  Mechanism  of  Labor  From  the  Standpoint  of  Comparative 
Anatomy,  With  a  Report  of  Cases  of  Dystocia  in  Wild 
Animals,  by  Edward  A.  Schumann,  M.D.  American  Jour- 
nal of  Obstetrics  and  Diseases  of  Women  and  Children,  Vol. 
LXIX,  No.  3,  19 1 4,  New  York. 

10.  Cirrhosis  of  the  Liver  in  Wild  Animals,  by  Herbert  Fox,  M.D. 

New  York  Medical  Journal,  December  19,  19 14. 

11.  The  Dynamics  of  the   Female   Pelvis;   Its   Evolution  and 

Architecture  with  Respect  to  Function,  by  Edward  A. 
Schumann,  M.D.  American  Journal  of  Obstetrics  and  Dis- 
eases of  Women  and  Children,  Vol.  LXXI,  No.  i,  1915, 
New  York. 

12.  Pneumonyssus  foxi,  Nov.  Sp.     An  Arachnid  Parasitic  in  the 

Lung  of  a  Monkey  (Macacus  rhesus),  by  Fred  D.  Weidman, 
M.D.  Journal  of  Parasitology,  September,  19 15,  Vol.  II,  pp. 
27-45,  Urbana,  Illinois. 

13.  Cytoleichus  penrosei,  A  New  Arachnid  Parasite  Found  in  the 

Diseased  Lungs  of  a  Prairie  Dog,  {Cynomys  ludovicianus) . 
Journal  of  Parasitology,  December,  1916,  Vol.  Ill,  pp.  82-89, 
Urbana,  Illinois.     Fred  D.  Weidman,  M.D. 

14.  A  Method  of  Obtaining  Duplicate  Reconstructions  from  the 

One  Series  of  Wax  Plates,  by  Fred  D.  Weidman,  M.D. 
New  York  Medical  Journal,  March  3,  191 7,  New  York. 

15.  Papers:  Read  at  the  Meeting  of  the  Pathological  Society  at 

the  Philadelphia  Zoological  Garden. 

Pancreatitis  in  Wild  Animals,  by  Herbert  Fox,  M.D. 

Report  of  an  Enzootic  of  Parasitic  Proven tricular  Worms 

{Spiroptera  incerta,  Smith)  of  Parrots,  with  Control  of 

Same,  by  Fred  D.  Weidman,  M.D, 

Coccidium  bigeminum.  Stiles,  in  Swift  Foxes   (habitat 

Western  U  S.),  by  Fred  D.  Weidman,  M.D. 

Distribution  of  Uncinaria  Among  the  Lower  Animals, 

by  Fred  D.  Weidman,  M.D. 

An  Arachnoid  {Pneumotuber  macaci,  Landois  and  Hoepke  ?) 

Parasitic  in  the  Lungs  of  a  Monkey  {Macacus  rhesus), 

by  Fred  D.  Weidman,  M.D. 

A  Note  Upon  the  Lesions  of  the  Female  Genitalia  in 

Wild  Animals,  by  Edward  A.  Shumann,  M.D. 


A  LIST  OF  PUBLICATIONS 

Amblyopia  in  a  Young  Monkey  (Macacus  nemestrinus) , 

by  H.  M.  Langdon,  M.D.  and  W.  B.  Cadawalder,  M.D. 

Remarks  on  Examinations  of  a  Series  of  Brains,  by  W.  B. 

Cadawalder,  M.D. 

Journal   of  Comparative  Pathology  and  Therapeutics, 

December,    1915,   Vol.   XXVIII.    Part    4,    pp.    298-336, 

Edinburgh,  Scotland. 

16.  Reversionary  Pseudobile  Canaliculi  Formation  in  the  Cirrhotic 

Liver  of  a  Vulpine  Phalanger,  by  Fred  D.  Weidman,  M.D. 
New  York  Medical  Journal,  March  9,  19 18,  New  York. 

17.  A  Contribution  to  the  Anatomy  and  Embryology  of  Cladorchis 

(Stichorchis)  Subtriquestrus,  Rudolphi,  18 14  (Fischoeder, 
1901),  by  Fred  D.  Weidman,  M.D.  Parasitology,  Vol.  X, 
No.  2,  January  22,  191 8,  Cambridge  University  Press, 
London,  England. 

18.  Nutritive  and  Blood  Changes  in  Rats  on  Cancer-Inhibiting 

and  Cancer-Stimulating  Diets,  by  E.  P.  Corson-White, 
M.D.  Pennsylvania  Medical  Journal,  March,  19 19,  Vol. 
XXII,  p.  348,  Athens,  Penna. 

19.  Pemphigus  in  an  Orang  Utan  Infested  with  Strongyloides 

(intestinalis  ?)  and  Dying  from  Advanced  Tuberculosis,  by 
Fred  D.  Weidman,  M.D.  Journal  of  Cutaneous  Diseases, 
March,  1919,  Vol.  XXXVII,  pp.  169-173,  Chicago,  111. 

20.  Arteriosclerosis  in  Wild  Animals,   by   Herbert   Fox,  M.D. 

American  Journal  of  Medical  Sciences,  June,  1920,  No.  6, 
Vol.  CLIX,  p.  821,  Philadelphia,  Penna. 

21.  Osteomalacia  in  Wild  Animals,  by  E.  P.  Corson-White,  M.D. 

Archives  of  Internal  Medicine,  November,  1922,  Vol.  30, 
pp.  620-628,  Chicago,  Illinois. 

22.  Osteitis   Deformans  in  Monkeys,  by  E.  P.  Corson-White, 

M.D.  Archives  of  Internal  Medicine,  December,  1922,  Vol. 
30,  pp.  790-796,  Chicago,  Illinois. 

23.  Certain  Dermatoses  of  Monkeys  and  an  Ape,  by  Fred  D. 

Weidman,  M.D.  Archives  of  Dermatology  and  Sy philology, 
March,  1923,  Vol.  7,  pp.  289-302,  Chicago,  Illinois. 

24.  Acute  Papular  and  Desquamative  Exanthem  in  an   Orang 

Utan,  by  Herbert  Fox,  M.D.,  and  Fred  D.  Weidman,  M.D. 
Archives  of  Dermatology  and  Syphilology,  April,  1923,  Vol.  7, 
pp.  462-464,  Chicago,  Illinois. 


(C-^    N.  MANCHESTER. 
'^■^  INDIANA, 


